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. Preface 

. An Introduction to the Human Body 

1. Overview 

2. Structural Organization of the Human Body 

3. Anatomical Terminology 

4. Medical Imaging 

. Body Tissues 

1. Types of Tissues 

2. Epithelial Tissue 

3. Connective Tissue Supports and Protects 

4. Muscle Tissue and Motion 

5. Nervous Tissue Mediates Perception and Response 

6. Tissue Injury and Aging 

. Integumentary System 

1. Layers of the Skin 

2. Accessory Structures of the Skin 

3. Diseases, Disorders, and Injuries of the Integumentary 
System 

. Bone 

1. The Functions of the Skeletal System 

2. Bone Classification 

3. Bone Structure 

4. Bone Formation and Development 

o. Fractures: Bone Repair 

. Axial Skeleton 

1. Divisions of the Skeletal System 

2. The Skull 

3. The Vertebral Column 

4. The Thoracic Cage 

. Appendicular Skeleton 

1. The Pectoral Girdle 


2. Bones of the Upper Limb 
3. The Pelvic Girdle and Pelvis 
4. Bones of the Lower Limb 
8. Articulations 
1. Classification of Joints 
2. Fibrous Joints 
3. Cartilaginous Joints 
4. Synovial Joints 
5. Types of Body Movements 
6. Anatomy of Selected Synovial Joints 
9. Skeletal Muscle 
1. Overview of Muscle Tissues 
2. Skeletal Muscle 
3. Types of Muscle Fibers 
4. Exercise and Muscle Performance 
5. Cardiac Muscle Tissue 
6. Smooth Muscle 
7. Development and Regeneration of Muscle Tissue 
8. Interactions of Skeletal Muscles, Their Fascicle 
Arrangement, and Their Lever Systems 
9. Naming Skeletal Muscles 
10. Axial Musculature 
1. Muscles of the Head, Neck, and Back 
2. Muscles of the Abdominal Wall and Thorax 
11. Appendicular Musculature 
1. Muscles of the Pectoral Girdle and Upper Limbs 
2. Muscles of the Pelvic Girdle and Lower Limbs 
12. Heart 
1. Heart Anatomy 
2. Cardiac Muscle and Electrical Activity 
3. Cardiac Cycle 
13. Blood Vessels 


14. 


15. 


16. 


17. 


18. 


19. 


20. 


21; 


22. 


1. Structure and Function of Blood Vessels 
2. Circulatory Pathways 
Blood 
1. An Overview of Blood 
2. Production of the Formed Elements 
3. Erythrocytes 
4. Leukocytes and Platelets 
Nervous Tissue 
1. Basic Structure and Function of the Nervous System 
2. Nervous ‘Tissue 
Central Nervous System 
1. Anatomy of the CNS 
2. Circulation and the Central Nervous System 
Peripheral Nervous System 
1. Nerves and ganglia 
Senses 
1. Sensory Perception 
Autonomic Nervous System 
1. Divisions of the ANS 
2. Central Control 
Respiratory System 
2. The Lungs 
Digestive System 
1. Overview of the Digestive System 


3. The Stomach 
4. The Small and Large Intestines 
5. Accessory Organs in Digestion: The Liver, Pancreas, and 
Gallbladder 
Lymphatic System 


23. Urinary System 


1. 
2. 
3: 


Gross Anatomy of Urine Transport 
Gross Anatomy of the Kidney 
Microscopic Anatomy of the Kidney 


24. Endocrine System 


1; 
. The Pituitary Gland and Hypothalamus 
. The Thyroid Gland 

. The Parathyroid Glands 

. The Adrenal Glands 

. The Pineal Gland 

. Gonadal and Placental Hormones 

. The Endocrine Pancreas 

9. 


CON MU BW NHN 


An Overview of the Endocrine System 


Organs with Secondary Endocrine Functions 


25. Reproductive System 


1. 
2. 


Male Anatomy 
Female Anatomy 


Preface 

Human Anatomy is designed for a semester-long course taken by life 
science and allied health students. The textbook is derived from OpenStax 
Human Anatomy and Physiology, and its coverage and organization were 
informed by hundreds of instructors who teach the course. Instructors can 
customize the book, adapting it to the approach that works best in their 
classroom. The artwork for this textbook is aimed focusing student learning 
through a powerful blend of traditional depictions and instructional 
innovations. Color is used sparingly, to emphasize the most important 
aspects of any given illustration. Significant use of micrographs from the 
University of Michigan complement the illustrations, and provide the 
students with a meaningful alternate depiction of each concept. Finally, 
enrichment elements provide relevance and deeper context for students, 
particularly in the areas of health, disease, and information relevant to their 
intended careers. 


Welcome to Human Anatomy, a resource designed for a semester-long 
course aimed at preparing undergraduate students for health-related 
programs. This book is derived from Human Anatomy and Physiology by 
OpenStax College. The source materials were created with several goals in 
mind: accessibility, customization, and student engagement—helping 
students reach high levels of academic scholarship. Instructors and students 
alike will find that this textbook offers a thorough introduction to the 
content in an accessible format. 


About OpenStax College 


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improve access to higher education for all. OpenStax College is an initiative 


of Rice University and is made possible through the generous support of 
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About OpenStax College’s Resources 


OpenStax College resources provide quality academic instruction. Three 
key features set our materials apart from others: 1) They can be easily 
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Human Anatomy and Physiology can be easily customized using our online 
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To broaden access and encourage community curation, Human Anatomy 
and Physiology is “open source” under a Creative Commons Attribution 
(CC BY) license. Members of the scientific community are invited to 


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Cost 


Our textbooks are available for free online, and in low-cost print and tablet 
editions. 


Contents 


01. 


02. 


03. 


04. 


05. 


06. 


07. 


08. 


09. 


Preface 

An Introduction to the Human Body 
Body Tissues 

Integumentary System 

Bone 

Axial Skeleton 

Appendicular Skeleton 
Articulations 


Skeletal Muscle 


10. Axial Musculature 

11. Appendicular Musculature 
12. Heart 

13. Blood Vessels 

14. Blood 

15. Nervous Tissue 

16. Central Nervous System 
17. Peripheral Nervous System 
18. Senses 

19. Autonomic Nervous System 
20. Respiratory System 

21. Digestive System 

22. Lymphatic System 

23. Urinary System 

24. Endocrine System 


25. Reproductive System 


About Our Team 


Customization for One-Semester Human Anatomy 


Marcos Gridi-Papp University of the Pacific 


Senior Contributors 


J. Gordon Betts Tyler Junior College 

Peter Desaix University of North Carolina at Chapel Hill 

Eddie Johnson Central Oregon Community College 

Jody E. Johnson Arapahoe Community College 

Oksana Korol Aims Community College 

Dean Kruse Portland Community College 

Brandon Poe Springfield Technical Community College 

James A. Wise Hampton University 

Mark Womble Youngstown State University 

Kelly A. Young California State University, Long Beach 
Advisor 


Robin J. Heyden 


Other Contributors 


Kim Aquarius Institute; Triton College 
Aaronson 

Lopamudra Augusta Technical College 

Agarwal 

Gary Allen Dalhousie University 

Robert i 

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Heather Southern Union State Community College 
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Ballard 

Matthew Eastern New Mexico University 

Barlow 

William Furman Universit 

Blaker 

Julie Bowers East Tennessee State University 

Emily Florida Southern College 

Bradshaw 


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Susan Caley 
Opsal 


Boyd 
Campbell 


Ann Caplea 


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Chapman 


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Crane 


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Culver 


Heather 
Cushman 


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Danzl-Tauer 


Jane Davis 


AnnMarie 
DelliPizzi 


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Pamela 
Dobbins 


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Health Sciences 


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Cornell College 


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Heinze 


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University of Maryland, Baltimore County 


Lihua Liang 


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Elisabeth 
Martin 


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Maxwell 


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McLaughlin 


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Mitchell 


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Natarajan 


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Nicotera 


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Niles 


Ikemefuna 
Nwosu 


Betsy Ott 


Ivan Paul 


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St. Bonaventure University 


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Parkland College; Lake Land College 


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St. Louis College of Pharmacy 


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Shobhana 
Natarajan 


Special Thanks 


University of Texas Medical Branch 
Oklahoma Baptist University 
Glendale Community College 
North Idaho College 


Bridgewater State University; Massasoit 
Community College 


Alcon Laboratories, Inc. 


OpenStax College wishes to thank the Regents of University of Michigan 

Medical School for the use of their extensive micrograph collection. Many 
of the UM micrographs that appear in Human Anatomy and Physiology are 
interactive WebScopes, which students can explore by zooming in and out. 


We also wish to thank the Open Learning Initiative at Carnegie Mellon 
University, with whom we shared and exchanged resources during the 
development of Human Anatomy and Physiology. 


Overview 
By the end of this section, you will be able to: 


¢ Compare and contrast anatomy and physiology, including their 
specializations and methods of study 
e Discuss the fundamental relationship between anatomy and physiology 


Human anatomy is the scientific study of the body’s structures. Some of 
these structures are very small and can only be observed and analyzed with 
the assistance of a microscope. Other larger structures can readily be seen, 
manipulated, measured, and weighed. The word “anatomy” comes from a 
Greek root that means “to cut apart.” Human anatomy was first studied by 
observing the exterior of the body and observing the wounds of soldiers and 
other injuries. Later, physicians were allowed to dissect bodies of the dead 
to augment their knowledge. When a body is dissected, its structures are cut 
apart in order to observe their physical attributes and their relationships to 
one another. Dissection is still used in medical schools, anatomy courses, 
and in pathology labs. In order to observe structures in living people, 
however, a number of imaging techniques have been developed. These 
techniques allow clinicians to visualize structures inside the living body 
such as a cancerous tumor or a fractured bone. 


Like most scientific disciplines, anatomy has areas of specialization. Gross 
anatomy is the study of the larger structures of the body, those visible 
without the aid of magnification ([link]a). Macro- means “large,” thus, 
gross anatomy is also referred to as macroscopic anatomy. In contrast, 
micro- means “small,” and microscopic anatomy is the study of structures 
that can be observed only with the use of a microscope or other 
magnification devices ({link]b). Microscopic anatomy includes cytology, 
the study of cells and histology, the study of tissues. As the technology of 
microscopes has advanced, anatomists have been able to observe smaller 
and smaller structures of the body, from slices of large structures like the 
heart, to the three-dimensional structures of large molecules in the body. 
Gross and Microscopic Anatomy 


(a) Gross anatomy considers large structures such as 
the brain. (b) Microscopic anatomy can deal with the 
same structures, though at a different scale. This is a 
micrograph of nerve cells from the brain. LM x 1600. 
(credit a: “WriterHound”/Wikimedia Commons; credit 
b: Micrograph provided by the Regents of University 
of Michigan Medical School © 2012) 


Anatomists take two general approaches to the study of the body’s 
structures: regional and systemic. Regional anatomy is the study of the 
interrelationships of all of the structures in a specific body region, such as 
the abdomen. Studying regional anatomy helps us appreciate the 
interrelationships of body structures, such as how muscles, nerves, blood 
vessels, and other structures work together to serve a particular body region. 
In contrast, systemic anatomy is the study of the structures that make up a 
discrete body system—that is, a group of structures that work together to 
perform a unique body function. For example, a systemic anatomical study 
of the muscular system would consider all of the skeletal muscles of the 
body. 


Whereas anatomy is about structure, physiology is about function. Human 
physiology is the scientific study of the chemistry and physics of the 
structures of the body and the ways in which they work together to support 
the functions of life. Much of the study of physiology centers on the body’s 
tendency toward homeostasis. Homeostasis is the state of steady internal 


conditions maintained by living things. The study of physiology certainly 
includes observation, both with the naked eye and with microscopes, as 
well as manipulations and measurements. However, current advances in 
physiology usually depend on carefully designed laboratory experiments 
that reveal the functions of the many structures and chemical compounds 
that make up the human body. 


Like anatomists, physiologists typically specialize in a particular branch of 
physiology. For example, neurophysiology is the study of the brain, spinal 
cord, and nerves and how these work together to perform functions as 
complex and diverse as vision, movement, and thinking. Physiologists may 
work from the organ level (exploring, for example, what different parts of 
the brain do) to the molecular level (such as exploring how an 
electrochemical signal travels along nerves). 


Form is closely related to function in all living things. For example, the thin 
flap of your eyelid can snap down to clear away dust particles and almost 
instantaneously slide back up to allow you to see again. At the microscopic 
level, the arrangement and function of the nerves and muscles that serve the 
eyelid allow for its quick action and retreat. At a smaller level of analysis, 
the function of these nerves and muscles likewise relies on the interactions 
of specific molecules and ions. Even the three-dimensional structure of 
certain molecules is essential to their function. 


Your study of anatomy and physiology will make more sense if you 
continually relate the form of the structures you are studying to their 
function. In fact, it can be somewhat frustrating to attempt to study anatomy 
without an understanding of the physiology that a body structure supports. 
Imagine, for example, trying to appreciate the unique arrangement of the 
bones of the human hand if you had no conception of the function of the 
hand. Fortunately, your understanding of how the human hand manipulates 
tools—from pens to cell phones—helps you appreciate the unique 
alignment of the thumb in opposition to the four fingers, making your hand 
a structure that allows you to pinch and grasp objects and type text 
messages. 


Chapter Review 


Human anatomy is the scientific study of the body’s structures. In the past, 
anatomy has primarily been studied via observing injuries, and later by the 
dissection of anatomical structures of cadavers, but in the past century, 
computer-assisted imaging techniques have allowed clinicians to look 
inside the living body. Human physiology is the scientific study of the 
chemistry and physics of the structures of the body. Physiology explains 
how the structures of the body work together to maintain life. It is difficult 
to study structure (anatomy) without knowledge of function (physiology). 
The two disciplines are typically studied together because form and 
function are closely related in all living things. 


Review Questions 


Exercise: 
Problem: 


Which of the following specialties might focus on studying all of the 
structures of the ankle and foot? 


a. microscopic anatomy 
b. muscle anatomy 

c. regional anatomy 

d. systemic anatomy 


Solution: 


C 
Exercise: 


Problem: 


A scientist wants to study how the body uses foods and fluids during a 
marathon run. This scientist is most likely a(n) 


a. exercise physiologist 
b. microscopic anatomist 


c. regional physiologist 
d. systemic anatomist 


Solution: 


A 


CRITICAL THINKING QUESTIONS 


Exercise: 


Problem: 
Name at least three reasons to study anatomy and physiology. 
Solution: 


An understanding of anatomy and physiology is essential for any 
career in the health professions. It can also help you make choices that 
promote your health, respond appropriately to signs of illness, make 
sense of health-related news, and help you in your roles as a parent, 
spouse, partner, friend, colleague, and caregiver. 


Exercise: 


Problem: 


For whom would an appreciation of the structural characteristics of the 
human heart come more easily: an alien who lands on Earth, abducts a 
human, and dissects his heart, or an anatomy and physiology student 
performing a dissection of the heart on her very first day of class? 
Why? 


Solution: 
A student would more readily appreciate the structures revealed in the 


dissection. Even though the student has not yet studied the workings of 
the heart and blood vessels in her class, she has experienced her heart 


beating every moment of her life, has probably felt her pulse, and 
likely has at least a basic understanding of the role of the heart in 
pumping blood throughout her body. This understanding of the heart’s 
function (physiology) would support her study of the heart’s form 
(anatomy). 


Glossary 


anatomy 
science that studies the form and composition of the body’s structures 


gross anatomy 
study of the larger structures of the body, typically with the unaided 
eye; also referred to macroscopic anatomy 


homeostasis 
steady state of body systems that living organisms maintain 


microscopic anatomy 
study of very small structures of the body using magnification 


physiology 
science that studies the chemistry, biochemistry, and physics of the 
body’s functions 


regional anatomy 
study of the structures that contribute to specific body regions 


systemic anatomy 
study of the structures that contribute to specific body systems 


Structural Organization of the Human Body 
By the end of this section, you will be able to: 


¢ Describe the structure of the human body in terms of six levels of 
organization 

e List the eleven organ systems of the human body and identify at least 
one organ and one major function of each 


Before you begin to study the different structures and functions of the 
human body, it is helpful to consider its basic architecture; that is, how its 
smallest parts are assembled into larger structures. It is convenient to 
consider the structures of the body in terms of fundamental levels of 
organization that increase in complexity: subatomic particles, atoms, 
molecules, organelles, cells, tissues, organs, organ systems, organisms and 
biosphere ({link]). 

Levels of Structural Organization of the Human Body 


Oxygen atom 


Chemical level 
Atoms bond to form 
molecules with three- 
dimensional structures. 


Jd Water molecule 


Smooth muscle cell 


Cellular level 
A variety of molecules 
combine to form the 
fluid and organelles 

of a body cell. 


Organelle 


Cell fluid 


Organelle 


Smooth muscle tissue 


Tissue level 
A community of similar 
cells form a body tissue. 


Bladder 


Organ level 
Two or more different tissues 
combine to form an organ. 


Skeletal 
muscle 


Urinary tract system 
Kidney 
Organ system level 
Ureter Two or more organs work 
closely together to perform 


he functi \f . 
Bladder the functions of a body system. 


A, Ureth ra 


1" 


Organismal level 
Many organ system work harmoniously 

together to perform the functions of an J 
independent organism. fo 


The organization of the body often is discussed in terms of six 
distinct levels of increasing complexity, from the smallest 
chemical building blocks to a unique human organism. 


The Levels of Organization 


To study the chemical level of organization, scientists consider the simplest 
building blocks of matter: subatomic particles, atoms and molecules. All 
matter in the universe is composed of one or more unique pure substances 
called elements, familiar examples of which are hydrogen, oxygen, carbon, 
nitrogen, calcium, and iron. The smallest unit of any of these pure 
substances (elements) is an atom. Atoms are made up of subatomic particles 
such as the proton, electron and neutron. Two or more atoms combine to 
form a molecule, such as the water molecules, proteins, and sugars found in 
living things. Molecules are the chemical building blocks of all body 
structures. 


A cell is the smallest independently functioning unit of a living organism. 
Even bacteria, which are extremely small, independently-living organisms, 
have a cellular structure. Each bacterium is a single cell. All living 
structures of human anatomy contain cells, and almost all functions of 
human physiology are performed in cells or are initiated by cells. 


A human cell typically consists of flexible membranes that enclose 
cytoplasm, a water-based cellular fluid together with a variety of tiny 
functioning units called organelles. In humans, as in all organisms, cells 
perform all functions of life. A tissue is a group of many similar cells 
(though sometimes composed of a few related types) that work together to 
perform a specific function. An organ is an anatomically distinct structure 
of the body composed of two or more tissue types. Each organ performs one 
or more specific physiological functions. An organ system is a group of 
organs that work together to perform major functions or meet physiological 
needs of the body. 


This book covers eleven distinct organ systems in the human body ([Link] 
and [link]). Assigning organs to organ systems can be imprecise since 
organs that “belong” to one system can also have functions integral to 
another system. In fact, most organs contribute to more than one system. 
Organ Systems of the Human Body 


Integumentary System Skeletal System 


¢ Encloses internal * Supports the body 
body structures ¢ Enables movement 
* Site of many (with muscular 
sensory receptors system) 
Muscular System Nervous System 
¢ Enables movement * Detects and 
(with skeletal system) processes sensory 
¢ Helps maintain body information 
temperature * Activates bodily 
responses 


Skeletal 
muscles 


Endocrine System Cardiovascular System 


Pituitary 

gland * Secretes hormones * Delivers oxygen 
. ¢ Regulates bodily and nutrients to 

Thyroid processes tissues 

gland * Equalizes 


temperature in 
the body 


Pancreas 


Adrenal 
glands 


Blood 
vessels 


Ovaries 


Organs that work together are grouped into organ 
systems. 


Organ Systems of the Human Body (continued) 


Lymphatic System 


¢ Returns fluid to 
blood 

¢ Defends against 
pathogens 


Digestive System 


¢ Processes food for 
use by the body 

« Removes wastes 
from undigested 
food 


Stomach 


Liver 


Gall 
bladder 


Large 
intestine 


Small 
intestine 


Male Reproductive 
System 


¢ Produces sex 
hormones and 
gametes 

* Delivers gametes 
to female 


Epididymis 


Respiratory System 


* Removes carbon 
dioxide from the 
body 

* Delivers oxygen 
to blood 


Nasal passage 


Trachea 


Lungs 


Urinary System 


* Controls water 
balance in the 
body 

« Removes wastes 
from blood and 


Kidneys excretes them 


Urinary 
bladder 


Female Reproductive 
System 
* Produces sex 
Mammary hormones 
glands and gametes 
* Supports embryo/ 
fetus until birth 
* Produces milk for 
: infant 
Ovaries 


Organs that work together are grouped into organ 
systems. 


The organism level is the highest level of organization. An organism is a 
living being that has a cellular structure and that can independently perform 
all physiologic functions necessary for life. In multicellular organisms, 
including humans, all cells, tissues, organs, and organ systems of the body 
work together to maintain the life and health of the organism. 


Chapter Review 


Life processes of the human body are maintained at several levels of 
structural organization. These include the chemical, cellular, tissue, organ, 
organ system, and the organism level. Higher levels of organization are 
built from lower levels. Therefore, molecules combine to form cells, cells 
combine to form tissues, tissues combine to form organs, organs combine to 
form organ systems, and organ systems combine to form organisms. 


Review Questions 


Exercise: 


Problem: 

The smallest independently functioning unit of an organism is a(n) 
a. cell 
b. molecule 


c. organ 
d. tissue 


Solution: 


A 
Exercise: 


Problem: 


A collection of similar tissues that performs a specific function is an 


a. organ 

b. organelle 

c. organism 

d. organ system 


Solution: 


A 
Exercise: 


Problem: 


The body system responsible for structural support and movement is 
the 


a. cardiovascular system 
b. endocrine system 

c. muscular system 

d. skeletal system 


Solution: 


D 


CRITICAL THINKING QUESTIONS 


Exercise: 


Problem:Name the six levels of organization of the human body. 
Solution: 


Chemical, cellular, tissue, organ, organ system, organism. 
Exercise: 


Problem: 


The female ovaries and the male testes are a part of which body 
system? Can these organs be members of more than one organ system? 
Why or why not? 


Solution: 


The female ovaries and the male testes are parts of the reproductive 
system. But they also secrete hormones, as does the endocrine system, 
therefore ovaries and testes function within both the endocrine and 
reproductive systems. 


Glossary 


cell 
smallest independently functioning unit of all organisms; in animals, a 
cell contains cytoplasm, composed of fluid and organelles 


organ 
functionally distinct structure composed of two or more types of 
tissues 


organ system 
group of organs that work together to carry out a particular function 


organism 
living being that has a cellular structure and that can independently 
perform all physiologic functions necessary for life 


tissue 
group of similar or closely related cells that act together to perform a 
specific function 


Anatomical Terminology 
By the end of this section, you will be able to: 


e Demonstrate the anatomical position 

e Describe the human body using directional and regional terms 

e Identify three planes most commonly used in the study of anatomy 

e Distinguish between the posterior (dorsal) and the anterior (ventral) 
body cavities, identifying their subdivisions and representative organs 
found in each 

e Describe serous membrane and explain its function 


Anatomists and health care providers use terminology that can be 
bewildering to the uninitiated. However, the purpose of this language is not 
to confuse, but rather to increase precision and reduce medical errors. For 
example, is a scar “above the wrist” located on the forearm two or three 
inches away from the hand? Or is it at the base of the hand? Is it on the 
palm-side or back-side? By using precise anatomical terminology, we 
eliminate ambiguity. Anatomical terms derive from ancient Greek and Latin 
words. Because these languages are no longer used in everyday 
conversation, the meaning of their words does not change. 


Anatomical terms are made up of roots, prefixes, and suffixes. The root of a 
term often refers to an organ, tissue, or condition, whereas the prefix or 
suffix often describes the root. For example, in the disorder hypertension, 
the prefix “hyper-” means “high” or “over,” and the root word “tension” 
refers to pressure, so the word “hypertension” refers to abnormally high 
blood pressure. 


Anatomical Position 


To further increase precision, anatomists standardize the way in which they 
view the body. Just as maps are normally oriented with north at the top, the 
standard body “map,” or anatomical position, is that of the body standing 
upright, with the feet at shoulder width and parallel, toes forward. The 
upper limbs are held out to each side, and the palms of the hands face 
forward as illustrated in [link]. Using this standard position reduces 
confusion. It does not matter how the body being described is oriented, the 


terms are used as if it is in anatomical position. For example, a scar in the 
“anterior (front) carpal (wrist) region” would be present on the palm side of 
the wrist. The term “anterior” would be used even if the hand were palm 
down on a table. 

Regions of the Human Body 


Frons or forehead (frontal). Oculus or eye 
(orbital or ocular) 


Cranium 
or skull 
(cranial) 


Bucca or cheek (buccal) Cephalon or head 


Shoulder (cephalic) 
(acromial) 


Facies 
or face 
(facial) 


Auris or ear (otic) 
Nasus or nose (nasal) 


Dorsum 
or back 


F Cervicis or neck 
Oris or mouth (oral) Cervicis or neck (cervical) 


Mentis or chin 


(dorsal) 


(mental) ; : Seaeun 
Axilla or armpit (thoracic) (brachial) 
(axillary) Mamma Olecranon 
. or breast or back 
bslhoral o (mammary) of elbow 
arm (brachial) ‘abdanner Trunk (olecranal) 
Antecubitis (abdominal) Lumbus 
or front of elbow or loin 
(antecubital) Umbilicus (lumbar) Upper 
or navel limb 
Antebrachium Sacrum 


(umbilical) 
or forearm 


(antebrachial) 
Carpus 
or wrist 
(carpal) 


Pollex Pelvis 


or thumb (pelvic) (manual) 
Palma or 

Inguen or groin Gluteus 
palm (palmar) ite inal) 9g sbatiock 
Digits (phalanges) (gluteal) 


ig 
or fingers (digital or 


phalangeal) aie or thigh 
Patella or (pubic) tone) 
ineecap Femur Poplit 
atellar) A opliteus or 
P ) . Naty back of knee 
Crus or (femoral) (popliteal) 
leg (crural) Sura 
or calf 
Tarsus (sural) 
or ankle 
(tarsal) Calcaneus or 
heel of foot 
Digits (phalanges) (calacaneal) 
or toes (digital or Pes or foot 
phalangeal) (pedal) Planta or sole 
~ of foot (plantar) 


(a) Anterior view 


Antebrachium 
or forearm 
(antebrachial) 


(sacral) 


Manus 
or hand 


(b) Posterior view 


The human body is shown in anatomical position in an (a) 
anterior view and a (b) posterior view. The regions of the body 
are labeled in boldface. 


A body that is lying down is described as either prone or supine. Prone 
describes a face-down orientation, and supine describes a face up 
orientation. These terms are sometimes used in describing the position of 
the body during specific physical examinations or surgical procedures. 


Regional Terms 


The human body’s numerous regions have specific terms to help increase 
precision (see [link]). Notice that the term “brachium” or “arm” is reserved 
for the “upper arm” and “antebrachium” or “forearm” is used rather than 
“lower arm.” Similarly, “femur” or “thigh” is correct, and “leg” or “crus” is 
reserved for the portion of the lower limb between the knee and the ankle. 
You will be able to describe the body’s regions using the terms from the 
figure. 


Directional Terms 


Certain directional anatomical terms appear throughout this and any other 
anatomy textbook ({link]). These terms are essential for describing the 
relative locations of different body structures. For instance, an anatomist 
might describe one band of tissue as “inferior to” another or a physician 
might describe a tumor as “superficial to” a deeper body structure. Commit 
these terms to memory to avoid confusion when you are studying or 
describing the locations of particular body parts. 


e Anterior (or ventral) Describes the front or direction toward the front 
of the body. The toes are anterior to the foot. 

¢ Posterior (or dorsal) Describes the back or direction toward the back 
of the body. The popliteus is posterior to the patella. 

e Superior (or cranial) describes a position above or higher than 
another part of the body proper. The orbits are superior to the oris. 

¢ Inferior (or caudal) describes a position below or lower than another 
part of the body proper; near or toward the tail (in humans, the coccyx, 
or lowest part of the spinal column). The pelvis is inferior to the 
abdomen. 

e Lateral describes the side or direction toward the side of the body. The 
thumb (pollex) is lateral to the digits. 

e Medial describes the middle or direction toward the middle of the 
body. The hallux is the medial toe. 

¢ Proximal describes a position in a limb that is nearer to the point of 
attachment or the trunk of the body. The brachium is proximal to the 
antebrachium. 


¢ Distal describes a position in a limb that is farther from the point of 
attachment or the trunk of the body. The crus is distal to the femur. 

e Superficial describes a position closer to the surface of the body. The 
skin is superficial to the bones. 

¢ Deep describes a position farther from the surface of the body. The 
brain is deep to the skull. 


Directional Terms Applied to the Human Body 


Superior 


Cranial 


-> Anterior or 
ventral 


Posterior +-7---- 
or dorsal 


Caudal 


Inferior 


Paired directional terms are shown as applied to 
the human body. 


Body Planes 


A section is a two-dimensional surface of a three-dimensional structure that 
has been cut. Modern medical imaging devices enable clinicians to obtain 
“virtual sections” of living bodies. We call these scans. Body sections and 
scans can be correctly interpreted, however, only if the viewer understands 
the plane along which the section was made. A plane is an imaginary two- 
dimensional surface that passes through the body. There are three planes 
commonly referred to in anatomy and medicine, as illustrated in [link]. 


e The sagittal plane is the plane that divides the body or an organ 
vertically into right and left sides. If this vertical plane runs directly 
down the middle of the body, it is called the midsagittal or median 
plane. If it divides the body into unequal right and left sides, it is called 
a parasagittal plane or less commonly a longitudinal section. 

e The frontal plane is the plane that divides the body or an organ into an 
anterior (front) portion and a posterior (rear) portion. The frontal plane 
is often referred to as a coronal plane. (“Corona” is Latin for “crown.”) 

e The transverse plane is the plane that divides the body or organ 
horizontally into upper and lower portions. Transverse planes produce 
images referred to as cross sections. 


Planes of the Body 


Frontal 
(coronal) 
plane 


Transverse 


The three planes most commonly used 
in anatomical and medical imaging 
are the sagittal, frontal (or coronal), 

and transverse plane. 


Body Cavities and Serous Membranes 


The body maintains its internal organization by means of membranes, 
sheaths, and other structures that separate compartments. The dorsal 
(posterior) cavity and the ventral (anterior) cavity are the largest body 
compartments ({link]). These cavities contain and protect delicate internal 
organs, and the ventral cavity allows for significant changes in the size and 
shape of the organs as they perform their functions. The lungs, heart, 
stomach, and intestines, for example, can expand and contract without 
distorting other tissues or disrupting the activity of nearby organs. 

Dorsal and Ventral Body Cavities 


Cranial 
cavity 


Vertebral 
cavity 


Thoracic cavity: 
Superior 
mediastinum 
Pleural cavity 
Pericardial 
cavity within 
the mediastinum 


Diaphragm 
Vertebral 


cavity Ventral body 


cavity 

(both thoracic and 
i i — 7 abdominopelvic 

Abdominal cavity Abdomino- | cavities) 

pelvic 

cavity 


Pelvic cavity 


Lateral view Anterior view 


The ventral cavity includes the thoracic and abdominopelvic 
cavities and their subdivisions. The dorsal cavity includes the 
cranial and spinal cavities. 


Subdivisions of the Posterior (Dorsal) and Anterior (Ventral) Cavities 


The posterior (dorsal) and anterior (ventral) cavities are each subdivided 
into smaller cavities. In the posterior (dorsal) cavity, the cranial cavity 
houses the brain, and the spinal cavity (or vertebral cavity) encloses the 
spinal cord. Just as the brain and spinal cord make up a continuous, 
uninterrupted structure, the cranial and spinal cavities that house them are 
also continuous. The brain and spinal cord are protected by the bones of the 
skull and vertebral column and by cerebrospinal fluid, a colorless fluid 
produced by the brain, which cushions the brain and spinal cord within the 
posterior (dorsal) cavity. 


The anterior (ventral) cavity has two main subdivisions: the thoracic cavity 
and the abdominopelvic cavity (see [link]). The thoracic cavity is the more 
superior subdivision of the anterior cavity, and it is enclosed by the rib cage. 
The thoracic cavity contains the lungs and the heart, which is located in the 
mediastinum. The diaphragm forms the floor of the thoracic cavity and 
separates it from the more inferior abdominopelvic cavity. The 
abdominopelvic cavity is the largest cavity in the body. Although no 
membrane physically divides the abdominopelvic cavity, it can be useful to 
distinguish between the abdominal cavity, the division that houses the 
digestive organs, and the pelvic cavity, the division that houses the organs 
of reproduction. 


Abdominal Regions and Quadrants 


To promote clear communication, for instance about the location of a 
patient’s abdominal pain or a suspicious mass, health care providers 
typically divide up the cavity into either nine regions or four quadrants 
({link]). 

Regions and Quadrants of the Peritoneal Cavity 


(a) Abdominopelvic regions (b) Abdominopelvic quandrants 


There are (a) nine abdominal regions and (b) four abdominal 
quadrants in the peritoneal cavity. 


The more detailed regional approach subdivides the cavity with one 
horizontal line immediately inferior to the ribs and one immediately 
superior to the pelvis, and two vertical lines drawn as if dropped from the 
midpoint of each clavicle (collarbone). There are nine resulting regions. 
The simpler quadrants approach, which is more commonly used in 
medicine, subdivides the cavity with one horizontal and one vertical line 
that intersect at the patient’s umbilicus (navel). 


Membranes of the Anterior (Ventral) Body Cavity 


A serous membrane (also referred to a serosa) is one of the thin 
membranes that cover the walls and organs in the thoracic and 
abdominopelvic cavities. The parietal layers of the membranes line the 
walls of the body cavity (pariet- refers to a cavity wall). The visceral layer 
of the membrane covers the organs (the viscera). Between the parietal and 
visceral layers is a very thin, fluid-filled serous space, or cavity ([link]). 
Serous Membrane 


Visceral 
pericardium 


Pericardial 
cavity 


Parietal 
pericardium 


\/— 


Air space 


Balloon 


Serous membrane lines the pericardial cavity 
and reflects back to cover the heart—much the 
same way that an underinflated balloon would 

form two layers surrounding a fist. 


There are three serous cavities and their associated membranes. The pleura 
is the serous membrane that encloses the pleural cavity; the pleural cavity 
surrounds the lungs. The pericardium is the serous membrane that encloses 
the pericardial cavity; the pericardial cavity surrounds the heart. The 
peritoneum is the serous membrane that encloses the peritoneal cavity; the 
peritoneal cavity surrounds several organs in the abdominopelvic cavity. 
The serous membranes form fluid-filled sacs, or cavities, that are meant to 
cushion and reduce friction on internal organs when they move, such as 
when the lungs inflate or the heart beats. Both the parietal and visceral 
serosa secrete the thin, slippery serous fluid located within the serous 
cavities. The pleural cavity reduces friction between the lungs and the body 
wall. Likewise, the pericardial cavity reduces friction between the heart and 
the wall of the pericardium. The peritoneal cavity reduces friction between 
the abdominal and pelvic organs and the body wall. Therefore, serous 
membranes provide additional protection to the viscera they enclose by 
reducing friction that could lead to inflammation of the organs. 


Chapter Review 


Ancient Greek and Latin words are used to build anatomical terms. A 
standard reference position for mapping the body’s structures is the normal 
anatomical position. Regions of the body are identified using terms such as 
“occipital” that are more precise than common words and phrases such as 
“the back of the head.” Directional terms such as anterior and posterior are 
essential for accurately describing the relative locations of body structures. 
Images of the body’s interior commonly align along one of three planes: the 
sagittal, frontal, or transverse. The body’s organs are organized in one of 
two main cavities—dorsal (also referred to posterior) and ventral (also 
referred to anterior)—which are further sub-divided according to the 
structures present in each area. The serous membranes have two layers— 
parietal and visceral—surrounding a fluid filled space. Serous membranes 
cover the lungs (pleural serosa), heart (pericardial serosa), and some 
abdominopelvic organs (peritoneal serosa). 


Review Chapter 


Exercise: 


Problem: 


What is the position of the body when it is in the “normal anatomical 
position?” 


a. The person is prone with upper limbs, including palms, touching 
sides and lower limbs touching at sides. 

b. The person is standing facing the observer, with upper limbs 
extended out at a ninety-degree angle from the torso and lower 
limbs in a wide stance with feet pointing laterally 

c. The person is supine with upper limbs, including palms, touching 
sides and lower limbs touching at sides. 

d. None of the above 


Solution: 


D 


Exercise: 


Problem: 


To make a banana split, you halve a banana into two long, thin, right 
and left sides along the 


a. coronal plane 

b. longitudinal plane 
c. midsagittal plane 
d. transverse plane 


Solution: 


C 


Exercise: 


Problem: The lumbar region is 


a. inferior to the gluteal region 

b. inferior to the umbilical region 
c. superior to the cervical region 
d. superior to the popliteal region 


Solution: 


D 


Exercise: 


Problem: The heart is within the 


a. Cranial cavity 

b. mediastinum 

c. posterior (dorsal) cavity 
d. All of the above 


Solution: 


B 


Critical Thinking Question 


Exercise: 


Problem: 


In which direction would an MRI scanner move to produce sequential 
images of the body in the frontal plane, and in which direction would 
an MRI scanner move to produce sequential images of the body in the 
sagittal plane? 


Solution: 


If the body were supine or prone, the MRI scanner would move from 
top to bottom to produce frontal sections, which would divide the body 
into anterior and posterior portions, as in “cutting” a deck of cards. 
Again, if the body were supine or prone, to produce sagittal sections, 
the scanner would move from left to right or from right to left to divide 
the body lengthwise into left and right portions. 


Exercise: 
Problem: 
If a bullet were to penetrate a lung, which three anterior thoracic body 


cavities would it enter, and which layer of the serous membrane would 
it encounter first? 


Solution: 


The bullet would enter the ventral, thoracic, and pleural cavities, and it 
would encounter the parietal layer of serous membrane first. 


Glossary 


abdominopelvic cavity 
division of the anterior (ventral) cavity that houses the abdominal and 
pelvic viscera 


anatomical position 
standard reference position used for describing locations and directions 
on the human body 


anterior 
describes the front or direction toward the front of the body; also 
referred to as ventral 


anterior cavity 
larger body cavity located anterior to the posterior (dorsal) body 
cavity; includes the serous membrane-lined pleural cavities for the 
lungs, pericardial cavity for the heart, and peritoneal cavity for the 
abdominal and pelvic organs; also referred to as ventral cavity 


caudal 
describes a position below or lower than another part of the body 
proper; near or toward the tail (in humans, the coccyx, or lowest part 
of the spinal column); also referred to as inferior 


cranial 
describes a position above or higher than another part of the body 
proper; also referred to as superior 


cranial cavity 
division of the posterior (dorsal) cavity that houses the brain 


deep 
describes a position farther from the surface of the body 


distal 
describes a position farther from the point of attachment or the trunk of 
the body 


dorsal 
describes the back or direction toward the back of the body; also 
referred to as posterior 


dorsal cavity 
posterior body cavity that houses the brain and spinal cord; also 
referred to the posterior body cavity 


frontal plane 
two-dimensional, vertical plane that divides the body or organ into 
anterior and posterior portions 


inferior 
describes a position below or lower than another part of the body 
proper; near or toward the tail (in humans, the coccyx, or lowest part 
of the spinal column); also referred to as caudal 


lateral 
describes the side or direction toward the side of the body 


medial 
describes the middle or direction toward the middle of the body 


pericardium 
sac that encloses the heart 


peritoneum 
serous membrane that lines the abdominopelvic cavity and covers the 
organs found there 


plane 
imaginary two-dimensional surface that passes through the body 


pleura 
serous membrane that lines the pleural cavity and covers the lungs 


posterior 


describes the back or direction toward the back of the body; also 
referred to as dorsal 


posterior cavity 
posterior body cavity that houses the brain and spinal cord; also 
referred to as dorsal cavity 


prone 
face down 


proximal 
describes a position nearer to the point of attachment or the trunk of 
the body 


sagittal plane 
two-dimensional, vertical plane that divides the body or organ into 
right and left sides 


section 
in anatomy, a single flat surface of a three-dimensional structure that 
has been cut through 


serous Membrane 
membrane that covers organs and reduces friction; also referred to as 
serosa 


serosa 
membrane that covers organs and reduces friction; also referred to as 
serous Membrane 


spinal cavity 
division of the dorsal cavity that houses the spinal cord; also referred 


to as vertebral cavity 


superficial 
describes a position nearer to the surface of the body 


superior 


describes a position above or higher than another part of the body 
proper; also referred to as cranial 


supine 
face up 


thoracic cavity 
division of the anterior (ventral) cavity that houses the heart, lungs, 
esophagus, and trachea 


transverse plane 
two-dimensional, horizontal plane that divides the body or organ into 
superior and inferior portions 


ventral 
describes the front or direction toward the front of the body; also 
referred to as anterior 


ventral cavity 
larger body cavity located anterior to the posterior (dorsal) body 
cavity; includes the serous membrane-lined pleural cavities for the 
lungs, pericardial cavity for the heart, and peritoneal cavity for the 
abdominal and pelvic organs; also referred to as anterior body cavity 


Medical Imaging 
By the end of this section, you will be able to: 


e Discuss the uses and drawbacks of X-ray imaging 
e Identify four modern medical imaging techniques and how they are 
used 


For thousands of years, fear of the dead and legal sanctions limited the 
ability of anatomists and physicians to study the internal structures of the 
human body. An inability to control bleeding, infection, and pain made 
surgeries infrequent, and those that were performed—such as wound 
suturing, amputations, tooth and tumor removals, skull drilling, and 
cesarean births—did not greatly advance knowledge about internal 
anatomy. Theories about the function of the body and about disease were 
therefore largely based on external observations and imagination. During 
the fourteenth and fifteenth centuries, however, the detailed anatomical 
drawings of Italian artist and anatomist Leonardo da Vinci and Flemish 
anatomist Andreas Vesalius were published, and interest in human anatomy 
began to increase. Medical schools began to teach anatomy using human 
dissection; although some resorted to grave robbing to obtain corpses. Laws 
were eventually passed that enabled students to dissect the corpses of 
criminals and those who donated their bodies for research. Still, it was not 
until the late nineteenth century that medical researchers discovered non- 
surgical methods to look inside the living body. 


X-Rays 


German physicist Wilhelm R6éntgen (1845-1923) was experimenting with 
electrical current when he discovered that a mysterious and invisible “ray” 
would pass through his flesh but leave an outline of his bones on a screen 
coated with a metal compound. In 1895, R6ntgen made the first durable 
record of the internal parts of a living human: an “X-ray” image (as it came 
to be called) of his wife’s hand. Scientists around the world quickly began 
their own experiments with X-rays, and by 1900, X-rays were widely used 
to detect a variety of injuries and diseases. In 1901, R6ntgen was awarded 
the first Nobel Prize for physics for his work in this field. 


The X-ray is a form of high energy electromagnetic radiation with a short 
wavelength capable of penetrating solids and ionizing gases. As they are 
used in medicine, X-rays are emitted from an X-ray machine and directed 
toward a specially treated metallic plate placed behind the patient’s body. 
The beam of radiation results in darkening of the X-ray plate. X-rays are 
slightly impeded by soft tissues, which show up as gray on the X-ray plate, 
whereas hard tissues, such as bone, largely block the rays, producing a 
light-toned “shadow.” Thus, X-rays are best used to visualize hard body 
structures such as teeth and bones ({link]). Like many forms of high energy 
radiation, however, X-rays are capable of damaging cells and initiating 
changes that can lead to cancer. This danger of excessive exposure to X- 
rays was not fully appreciated for many years after their widespread use. 
X-Ray of a Hand 


High energy 
electromagnetic radiation 
allows the internal 
structures of the body, such 
as bones, to be seen in X- 
rays like these. (credit: 
Trace Meek/flickr) 


Refinements and enhancements of X-ray techniques have continued 
throughout the twentieth and twenty-first centuries. Although often 
supplanted by more sophisticated imaging techniques, the X-ray remains a 
“workhorse” in medical imaging, especially for viewing fractures and for 
dentistry. The disadvantage of irradiation to the patient and the operator is 
now attenuated by proper shielding and by limiting exposure. 


Modern Medical Imaging 


X-rays can depict a two-dimensional image of a body region, and only from 
a single angle. In contrast, more recent medical imaging technologies 
produce data that is integrated and analyzed by computers to produce three- 
dimensional images or images that reveal aspects of body functioning. 


Computed Tomography 


Tomography refers to imaging by sections. Computed tomography (CT) 
is a noninvasive imaging technique that uses computers to analyze several 
cross-sectional X-rays in order to reveal minute details about structures in 
the body ({link]a). The technique was invented in the 1970s and is based on 
the principle that, as X-rays pass through the body, they are absorbed or 
reflected at different levels. In the technique, a patient lies on a motorized 
platform while a computerized axial tomography (CAT) scanner rotates 360 
degrees around the patient, taking X-ray images. A computer combines 
these images into a two-dimensional view of the scanned area, or “slice.” 
Medical Imaging Techniques 


(a) The results of a CT scan of the head are shown as 
successive transverse sections. (b) An MRI machine 
generates a magnetic field around a patient. (c) PET 
scans use radiopharmaceuticals to create images of 
active blood flow and physiologic activity of the organ 
or organs being targeted. (d) Ultrasound technology is 
used to monitor pregnancies because it is the least 
invasive of imaging techniques and uses no 
electromagnetic radiation. (credit a: Akira 
Ohgaki/flickr; credit b: “Digital Cate”/flickr; credit c: 
“Raziel”/Wikimedia Commons; credit d: 
“Tsis”/Wikimedia Commons) 


Since 1970, the development of more powerful computers and more 
sophisticated software has made CT scanning routine for many types of 


diagnostic evaluations. It is especially useful for soft tissue scanning, such 
as of the brain and the thoracic and abdominal viscera. Its level of detail is 
so precise that it can allow physicians to measure the size of a mass down to 
a millimeter. The main disadvantage of CT scanning is that it exposes 
patients to a dose of radiation many times higher than that of X-rays. In 
fact, children who undergo CT scans are at increased risk of developing 
cancer, as are adults who have multiple CT scans. 


Note: 
wae 


— : 
mess OPenstax COLLEGE 


A CT or CAT scan relies on a circling scanner that revolves around the 
patient’s body. Watch this video to learn more about CT and CAT scans. 
What type of radiation does a CT scanner use? 


Magnetic Resonance Imaging 


Magnetic resonance imaging (MRI) is a noninvasive medical imaging 
technique based on a phenomenon of nuclear physics discovered in the 
1930s, in which matter exposed to magnetic fields and radio waves was 
found to emit radio signals. In 1970, a physician and researcher named 
Raymond Damadian noticed that malignant (cancerous) tissue gave off 
different signals than normal body tissue. He applied for a patent for the 
first MRI scanning device, which was in use clinically by the early 1980s. 
The early MRI scanners were crude, but advances in digital computing and 
electronics led to their advancement over any other technique for precise 
imaging, especially to discover tumors. MRI also has the major advantage 
of not exposing patients to radiation. 


Drawbacks of MRI scans include their much higher cost, and patient 
discomfort with the procedure. The MRI scanner subjects the patient to 
such powerful electromagnets that the scan room must be shielded. The 
patient must be enclosed in a metal tube-like device for the duration of the 
scan (see [link]b), sometimes as long as thirty minutes, which can be 
uncomfortable and impractical for ill patients. The device is also so noisy 
that, even with earplugs, patients can become anxious or even fearful. 
These problems have been overcome somewhat with the development of 
“open” MRI scanning, which does not require the patient to be entirely 
enclosed in the metal tube. Patients with iron-containing metallic implants 
(internal sutures, some prosthetic devices, and so on) cannot undergo MRI 
scanning because it can dislodge these implants. 


Functional MRIs ({MRIs), which detect the concentration of blood flow in 
certain parts of the body, are increasingly being used to study the activity in 
parts of the brain during various body activities. This has helped scientists 
learn more about the locations of different brain functions and more about 
brain abnormalities and diseases. 


Note: 


zis 


A patient undergoing an MRI is surrounded by a tube-shaped scanner. 
Watch this video to learn more about MRIs. What is the function of 
magnets in an MRI? 


Positron Emission Tomography 


Positron emission tomography (PET) is a medical imaging technique 
involving the use of so-called radiopharmaceuticals, substances that emit 
radiation that is short-lived and therefore relatively safe to administer to the 
body. Although the first PET scanner was introduced in 1961, it took 15 
more years before radiopharmaceuticals were combined with the technique 
and revolutionized its potential. The main advantage is that PET (see 
[link]c) can illustrate physiologic activity—including nutrient metabolism 
and blood flow—of the organ or organs being targeted, whereas CT and 
MRI scans can only show static images. PET is widely used to diagnose a 
multitude of conditions, such as heart disease, the spread of cancer, certain 
forms of infection, brain abnormalities, bone disease, and thyroid disease. 


PET relies on radioactive substances administered several minutes before 
the scan. Watch this video to learn more about PET. How is PET used in 
chemotherapy? 


Ultrasonography 


Ultrasonography is an imaging technique that uses the transmission of 
high-frequency sound waves into the body to generate an echo signal that is 
converted by a computer into a real-time image of anatomy and physiology 
(see [link]d). Ultrasonography is the least invasive of all imaging 
techniques, and it is therefore used more freely in sensitive situations such 
as pregnancy. The technology was first developed in the 1940s and 1950s. 
Ultrasonography is used to study heart function, blood flow in the neck or 
extremities, certain conditions such as gallbladder disease, and fetal growth 


and development. The main disadvantages of ultrasonography are that the 
image quality is heavily operator-dependent and that it is unable to 
penetrate bone and gas. 


Chapter Review 


Detailed anatomical drawings of the human body first became available in 
the fifteenth and sixteenth centuries; however, it was not until the end of the 
nineteenth century, and the discovery of X-rays, that anatomists and 
physicians discovered non-surgical methods to look inside a living body. 
Since then, many other techniques, including CT scans, MRI scans, PET 
scans, and ultrasonography, have been developed, providing more accurate 
and detailed views of the form and function of the human body. 


Interactive Link Questions 


Exercise: 
Problem: 
A CT or CAT scan relies on a circling scanner that revolves around the 


patient’s body. Watch this video to learn more about CT and CAT 
scans. What type of radiation does a CT scanner use? 


Solution: 


X-rays. 
Exercise: 
Problem: 
A patient undergoing an MRI is surrounded by a tube-shaped scanner. 


Watch this video to learn more about MRIs. What is the function of 
magnets in an MRI? 


Solution: 


The magnets induce tissue to emit radio signals that can show 
differences between different types of tissue. 


Exercise: 
Problem: 
PET relies on radioactive substances administered several minutes 


before the scan. Watch this video to learn more about PET. How is 
PET used in chemotherapy? 


Solution: 


PET scans can indicate how patients are responding to chemotherapy. 


Review Questions 


Exercise: 
Problem: 


In 1901, Wilhelm Réntgen was the first person to win the Nobel Prize 
for physics. For what discovery did he win? 


a. nuclear physics 

b. radiopharmaceuticals 

c. the link between radiation and cancer 
d. X-rays 


Solution: 


D 
Exercise: 


Problem: 


Which of the following imaging techniques would be best to use to 
study the uptake of nutrients by rapidly multiplying cancer cells? 


a. CT 

b. MRI 

c. PET 

d. ultrasonography 


Solution: 


C 
Exercise: 
Problem: 


Which of the following imaging studies can be used most safely during 
pregnancy? 


a. CT scans 
b. PET scans 
c. ultrasounds 
d. X-rays 


Solution: 


C 


Exercise: 


Problem: What are two major disadvantages of MRI scans? 


a. release of radiation and poor quality images 

b. high cost and the need for shielding from the magnetic signals 

c. can only view metabolically active tissues and inadequate 
availability of equipment 

d. release of radiation and the need for a patient to be confined to 
metal tube for up to 30 minutes 


Solution: 


Critical Thinking Questions 


Exercise: 
Problem: 


Which medical imaging technique is most dangerous to use repeatedly, 
and why? 


Solution: 
CT scanning subjects patients to much higher levels of radiation than 
X-rays, and should not be performed repeatedly. 
Exercise: 
Problem: 


Explain why ultrasound imaging is the technique of choice for 
studying fetal growth and development. 


Solution: 


Ultrasonography does not expose a mother or fetus to radiation, to 
radiopharmaceuticals, or to magnetic fields. At this time, there are no 
known medical risks of ultrasonography. 


Glossary 


computed tomography (CT) 
medical imaging technique in which a computer-enhanced cross- 
sectional X-ray image is obtained 


magnetic resonance imaging (MRI) 
medical imaging technique in which a device generates a magnetic 
field to obtain detailed sectional images of the internal structures of the 


body 


positron emission tomography (PET) 
medical imaging technique in which radiopharmaceuticals are traced 
to reveal metabolic and physiological functions in tissues 


ultrasonography 
application of ultrasonic waves to visualize subcutaneous body 
structures such as tendons and organs 


X-ray 
form of high energy electromagnetic radiation with a short wavelength 
capable of penetrating solids and ionizing gases; used in medicine as a 
diagnostic aid to visualize body structures such as bones 


Types of Tissues 
By the end of this section, you will be able to: 


e Identify the four main tissue types 

e Discuss the functions of each tissue type 

e Relate the structure of each tissue type to their function 
e Discuss the embryonic origin of tissue 

e Identify the three major germ layers 

e Identify the main types of tissue membranes 


The term tissue is used to describe a group of cells found together in the 
body. The cells within a tissue share a common embryonic origin. 
Microscopic observation reveals that the cells in a tissue share 
morphological features and are arranged in an orderly pattern that achieves 
the tissue’s functions. From the evolutionary perspective, tissues appear in 
more complex organisms. For example, multicellular protists, ancient 
eukaryotes, do not have cells organized into tissues. 


Although there are many types of cells in the human body, they are 
organized into four broad categories of tissues: epithelial, connective, 
muscle, and nervous. Each of these categories is characterized by specific 
functions that contribute to the overall health and maintenance of the body. 
A disruption of the structure is a sign of injury or disease. Such changes can 
be detected through histology, the microscopic study of tissue appearance, 
organization, and function. 


The Four Types of Tissues 


Epithelial tissue, also referred to as epithelium, refers to the sheets of cells 
that cover exterior surfaces of the body, lines internal cavities and 
passageways, and forms certain glands. Connective tissue, as its name 
implies, binds the cells and organs of the body together and functions in the 
protection, support, and integration of all parts of the body. Muscle tissue is 
excitable, responding to stimulation and contracting to provide movement, 
and occurs as three major types: skeletal (voluntary) muscle, smooth 
muscle, and cardiac muscle in the heart. Nervous tissue is also excitable, 


allowing the propagation of electrochemical signals in the form of nerve 
impulses that communicate between different regions of the body ([link]). 


The next level of organization is the organ, where several types of tissues 
come together to form a working unit. Just as knowing the structure and 
function of cells helps you in your study of tissues, knowledge of tissues 
will help you understand how organs function. The epithelial and 
connective tissues are discussed in detail in this chapter. Muscle and 
nervous tissues will be discussed only briefly in this chapter. 

Four Types of Tissue: Body 


Nervous tissue 
Brain 
Spinal cord 
Nerves 


Muscle tissue 
Cardiac muscle 
Smooth muscle 
Skeletal muscle 


Epithelial tissue 
Lining of Gl tract organs 
and other hollow organs 
Skin surface (epidermis) 


Connective tissue 
Fat and other soft 
padding tissue 

Bone 
Tendon 


The four types of tissues are exemplified in nervous 
tissue, stratified squamous epithelial tissue, cardiac 
muscle tissue, and connective tissue in small intestine. 


Clockwise from nervous tissue, LM x 872, LM x 282, 

LM x 460, LM x 800. (Micrographs provided by the 

Regents of University of Michigan Medical School © 
2012) 


Embryonic Origin of Tissues 


The zygote, or fertilized egg, is a single cell formed by the fusion of an egg 
and sperm. After fertilization the zygote gives rise to rapid mitotic cycles, 
generating many cells to form the embryo. The first embryonic cells 
generated have the ability to differentiate into any type of cell in the body 
and, as such, are called totipotent, meaning each has the capacity to divide, 
differentiate, and develop into a new organism. As cell proliferation 
progresses, three major cell lineages are established within the embryo. As 
explained in a later chapter, each of these lineages of embryonic cells forms 
the distinct germ layers from which all the tissues and organs of the human 
body eventually form. Each germ layer is identified by its relative position: 
ectoderm (ecto- = “outer”), mesoderm (meso- = “middle”), and endoderm 
(endo- = “inner’’). [link] shows the types of tissues and organs associated 
with the each of the three germ layers. Note that epithelial tissue originates 
in all three layers, whereas nervous tissue derives primarily from the 
ectoderm and muscle tissue from mesoderm. 

Embryonic Origin of Tissues and Major Organs 


Ectoderm Epidermis, glands on skin, some cranial bones, pituitary and adrenal medulla, the nervous 
system, the mouth between cheek and gums, the anus 
= ue, ANZ 1Q( | a 
= is S P= y =. ] : y/ | . = Nos / 
Skin cells Neurons Pigment cell 
Mesoderm Connective tissues proper, bone, cartilage, blood, endothelium of blood vessels, muscle, 
synovial membranes, serous membranes lining body cavities, kidneys, lining of gonads 
_——— 
—— = — = 
— 
be _<<ff = = 
Cardiac Skeletal Tubule cell Red blood Smooth 
muscle muscle of kidney cells muscle 
Endoderm Lining of airways and digestive system except the mouth and distal part of digestive system 
(rectum and anal canal); glands (digestive glands, endocrine glands, adrenal cortex) 
Lung cell Thyroid cell Pancreatic cell 


= = : 
mmm §~OPENStax COLLEGE 


View this slideshow to learn more about stem cells. How do somatic stem 
cells differ from embryonic stem cells? 


Tissue Membranes 


A tissue membrane is a thin layer or sheet of cells that covers the outside 
of the body (for example, skin), the organs (for example, pericardium), 
internal passageways that lead to the exterior of the body (for example, 
abdominal mesenteries), and the lining of the moveable joint cavities. There 
are two basic types of tissue membranes: connective tissue and epithelial 
membranes ((link]). 

Tissue Membranes 


Mucous membranes line the 
digestive, respiratory, urinary, 
and reproductive tracts. They 
are coated with the secretions 
of mucous glands. 


Serous membranes line body 
cavities closed to the exterior 
of the body: the peritoneal, 
pleural, and pericardial 
cavities. 


Cutaneous membrane, or the 
skin, covers the body surface. 


Synovial membranes line joint 
cavities and produce the fluid 


within the joint. 


The two broad categories of tissue membranes 
in the body are (1) connective tissue 
membranes, which include synovial 

membranes, and (2) epithelial membranes, 
which include mucous membranes, serous 
membranes, and the cutaneous membrane, in 
other words, the skin. 


Connective Tissue Membranes 


The connective tissue membrane is formed solely from connective tissue. 
These membranes encapsulate organs, such as the kidneys, and line our 
movable joints. A synovial membrane is a type of connective tissue 
membrane that lines the cavity of a freely movable joint. For example, 
synovial membranes surround the joints of the shoulder, elbow, and knee. 
Fibroblasts in the inner layer of the synovial membrane release hyaluronan 
into the joint cavity. The hyaluronan effectively traps available water to 
form the synovial fluid, a natural lubricant that enables the bones of a joint 
to move freely against one another without much friction. This synovial 
fluid readily exchanges water and nutrients with blood, as do all body 
fluids. 


Epithelial Membranes 


The epithelial membrane is composed of epithelium attached to a layer of 
connective tissue, for example, your skin. The mucous membrane is also a 
composite of connective and epithelial tissues. Sometimes called mucosae, 
these epithelial membranes line the body cavities and hollow passageways 
that open to the external environment, and include the digestive, respiratory, 
excretory, and reproductive tracts. Mucous, produced by the epithelial 
exocrine glands, covers the epithelial layer. The underlying connective 
tissue, called the lamina propria (literally “own layer”), help support the 
fragile epithelial layer. 


A serous membrane is an epithelial membrane composed of mesodermally 
derived epithelium called the mesothelium that is supported by connective 
tissue. These membranes line the coelomic cavities of the body, that is, 
those cavities that do not open to the outside, and they cover the organs 
located within those cavities. They are essentially membranous bags, with 
mesothelium lining the inside and connective tissue on the outside. Serous 
fluid secreted by the cells of the thin squamous mesothelium lubricates the 
membrane and reduces abrasion and friction between organs. Serous 
membranes are identified according locations. Three serous membranes line 
the thoracic cavity; the two pleura that cover the lungs and the pericardium 


that covers the heart. A fourth, the peritoneum, is the serous membrane in 
the abdominal cavity that covers abdominal organs and forms double sheets 
of mesenteries that suspend many of the digestive organs. 


The skin is an epithelial membrane also called the cutaneous membrane. It 
is a stratified squamous epithelial membrane resting on top of connective 
tissue. The apical surface of this membrane is exposed to the external 
environment and is covered with dead, keratinized cells that help protect the 
body from desiccation and pathogens. 


Chapter Review 


The human body contains more than 200 types of cells that can all be 
classified into four types of tissues: epithelial, connective, muscle, and 
nervous. Epithelial tissues act as coverings controlling the movement of 
materials across the surface. Connective tissue integrates the various parts 
of the body and provides support and protection to organs. Muscle tissue 
allows the body to move. Nervous tissues propagate information. 


The study of the shape and arrangement of cells in tissue is called histology. 
All cells and tissues in the body derive from three germ layers in the 
embryo: the ectoderm, mesoderm, and endoderm. 


Different types of tissues form membranes that enclose organs, provide a 
friction-free interaction between organs, and keep organs together. Synovial 
membranes are connective tissue membranes that protect and line the joints. 
Epithelial membranes are formed from epithelial tissue attached to a layer 
of connective tissue. There are three types of epithelial membranes: 
mucous, which contain glands; serous, which secrete fluid; and cutaneous 
which makes up the skin. 


Interactive Link Questions 


Exercise: 


Problem: 


View this slideshow to learn more about stem cells. How do somatic 
stem cells differ from embryonic stem cells? 


Solution: 


Most somatic stem cells give rise to only a few cell types. 


Review Questions 


Exercise: 


Problem: Which of the following is not a type of tissue? 


a. muscle 

b. nervous 

c. embryonic 
d. epithelial 


Solution: 


c 
Exercise: 
Problem: 


The process by which a less specialized cell matures into a more 
specialized cell is called 


a. differentiation 
b. maturation 

c. modification 
d. specialization 


Solution: 


A 
Exercise: 


Problem: 
Differentiated cells in a developing embryo derive from 


a. endothelium, mesothelium, and epithelium 

b. ectoderm, mesoderm, and endoderm 

c. connective tissue, epithelial tissue, and muscle tissue 
d. epidermis, mesoderm, and endothelium 


Solution: 


B 
Exercise: 


Problem: 


Which of the following lines the body cavities exposed to the external 
environment? 


a. mesothelium 
b. lamina propria 


c. mesenteries 
d. mucosa 


Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: 


Identify the four types of tissue in the body, and describe the major 
functions of each tissue. 


Solution: 


The four types of tissue in the body are epithelial, connective, muscle, 
and nervous. Epithelial tissue is made of layers of cells that cover the 
surfaces of the body that come into contact with the exterior world, 
line internal cavities, and form glands. Connective tissue binds the 
cells and organs of the body together and performs many functions, 
especially in the protection, support, and integration of the body. 
Muscle tissue, which responds to stimulation and contracts to provide 
movement, is divided into three major types: skeletal (voluntary) 
muscles, smooth muscles, and the cardiac muscle in the heart. Nervous 
tissue allows the body to receive signals and transmit information as 
electric impulses from one region of the body to another. 


Exercise: 


Problem: 


The zygote is described as totipotent because it ultimately gives rise to 
all the cells in your body including the highly specialized cells of your 
nervous system. Describe this transition, discussing the steps and 
processes that lead to these specialized cells. 


Solution: 


The zygote divides into many cells. As these cells become specialized, 
they lose their ability to differentiate into all tissues. At first they form 
the three primary germ layers. Following the cells of the ectodermal 
germ layer, they too become more restricted in what they can form. 
Ultimately, some of these ectodermal cells become further restricted 
and differentiate in to nerve cells. 


Exercise: 


Problem: What is the function of synovial membranes? 
Solution: 


Synovial membranes are a type of connective tissue membrane that 
supports mobility in joints. The membrane lines the joint cavity and 
contains fibroblasts that produce hyaluronan, which leads to the 
production of synovial fluid, a natural lubricant that enables the bones 
of a joint to move freely against one another. 


Glossary 


connective tissue 
type of tissue that serves to hold in place, connect, and integrate the 
body’s organs and systems 


connective tissue membrane 
connective tissue that encapsulates organs and lines movable joints 


cutaneous membrane 
skin; epithelial tissue made up of a stratified squamous epithelial cells 
that cover the outside of the body 


ectoderm 
outermost embryonic germ layer from which the epidermis and the 
nervous tissue derive 


endoderm 
innermost embryonic germ layer from which most of the digestive 
system and lower respiratory system derive 


epithelial membrane 
epithelium attached to a layer of connective tissue 


epithelial tissue 


type of tissue that serves primarily as a covering or lining of body 
parts, protecting the body; it also functions in absorption, transport, 
and secretion 


histology 
microscopic study of tissue architecture, organization, and function 


lamina propria 
areolar connective tissue underlying a mucous membrane 


mesoderm 
middle embryonic germ layer from which connective tissue, muscle 
tissue, and some epithelial tissue derive 


mucous membrane 
tissue membrane that is covered by protective mucous and lines tissue 
exposed to the outside environment 


muscle tissue 
type of tissue that is capable of contracting and generating tension in 
response to stimulation; produces movement. 


nervous tissue 
type of tissue that is capable of sending and receiving impulses 
through electrochemical signals. 


serous membrane 
type of tissue membrane that lines body cavities and lubricates them 
with serous fluid 


synovial membrane 
connective tissue membrane that lines the cavities of freely movable 
joints, producing synovial fluid for lubrication 


tissue 
group of cells that are similar in form and perform related functions 


tissue membrane 


thin layer or sheet of cells that covers the outside of the body, organs, 
and internal cavities 


totipotent 
embryonic cells that have the ability to differentiate into any type of 
cell and organ in the body 


Epithelial Tissue 
By the end of this section, you will be able to: 


e Explain the structure and function of epithelial tissue 

e Distinguish between tight junctions, anchoring junctions, and gap 
junctions 

e Distinguish between simple epithelia and stratified epithelia, as well as 
between squamous, cuboidal, and columnar epithelia 

e Describe the structure and function of endocrine and exocrine glands 
and their respective secretions 


Most epithelial tissues are essentially large sheets of cells covering all the 
surfaces of the body exposed to the outside world and lining the outside of 
organs. Epithelium also forms much of the glandular tissue of the body. 
Skin is not the only area of the body exposed to the outside. Other areas 
include the airways, the digestive tract, as well as the urinary and 
reproductive systems, all of which are lined by an epithelium. Hollow 
organs and body cavities that do not connect to the exterior of the body, 
which includes, blood vessels and serous membranes, are lined by 
endothelium (plural = endothelia), which is a type of epithelium. 


Epithelial cells derive from all three major embryonic layers. The epithelia 
lining the skin, parts of the mouth and nose, and the anus develop from the 
ectoderm. Cells lining the airways and most of the digestive system 
originate in the endoderm. The epithelium that lines vessels in the 
lymphatic and cardiovascular system derives from the mesoderm and is 
called an endothelium. 


All epithelia share some important structural and functional features. This 
tissue is highly cellular, with little or no extracellular material present 
between cells. Adjoining cells form a specialized intercellular connection 
between their cell membranes called a cell junction. The epithelial cells 
exhibit polarity with differences in structure and function between the 
exposed or apical facing surface of the cell and the basal surface close to 
the underlying body structures. The basal lamina, a mixture of 
glycoproteins and collagen, provides an attachment site for the epithelium, 
separating it from underlying connective tissue. The basal lamina attaches 


to a reticular lamina, which is secreted by the underlying connective 
tissue, forming a basement membrane that helps hold it all together. 


Epithelial tissues are nearly completely avascular. For instance, no blood 
vessels cross the basement membrane to enter the tissue, and nutrients must 
come by diffusion or absorption from underlying tissues or the surface. 
Many epithelial tissues are capable of rapidly replacing damaged and dead 
cells. Sloughing off of damaged or dead cells is a characteristic of surface 
epithelium and allows our airways and digestive tracts to rapidly replace 
damaged cells with new cells. 


Generalized Functions of Epithelial Tissue 


Epithelial tissues provide the body’s first line of protection from physical, 
chemical, and biological wear and tear. The cells of an epithelium act as 
gatekeepers of the body controlling permeability and allowing selective 
transfer of materials across a physical barrier. All substances that enter the 
body must cross an epithelium. Some epithelia often include structural 
features that allow the selective transport of molecules and ions across their 
cell membranes. 


Many epithelial cells are capable of secretion and release mucous and 
specific chemical compounds onto their apical surfaces. The epithelium of 
the small intestine releases digestive enzymes, for example. Cells lining the 
respiratory tract secrete mucous that traps incoming microorganisms and 
particles. A glandular epithelium contains many secretory cells. 


The Epithelial Cell 


Epithelial cells are typically characterized by the polarized distribution of 
organelles and membrane-bound proteins between their basal and apical 
surfaces. Particular structures found in some epithelial cells are an 
adaptation to specific functions. Certain organelles are segregated to the 
basal sides, whereas other organelles and extensions, such as cilia, when 
present, are on the apical surface. 


Cilia are microscopic extensions of the apical cell membrane that are 
supported by microtubules. They beat in unison and move fluids as well as 
trapped particles. Ciliated epithelium lines the ventricles of the brain where 
it helps circulate the cerebrospinal fluid. The ciliated epithelium of your 
airway forms a mucociliary escalator that sweeps particles of dust and 
pathogens trapped in the secreted mucous toward the throat. It is called an 
escalator because it continuously pushes mucous with trapped particles 
upward. In contrast, nasal cilia sweep the mucous blanket down towards 
your throat. In both cases, the transported materials are usually swallowed, 
and end up in the acidic environment of your stomach. 


Cell to Cell Junctions 


Cells of epithelia are closely connected and are not separated by 
intracellular material. Three basic types of connections allow varying 
degrees of interaction between the cells: tight junctions, anchoring 
junctions, and gap junctions ([link]). 

Types of Cell Junctions 


Tight junction 


plasma 
membranes 


Strands of <@ 


transmembrane 
proteins 


Intercellullar 


Anchoring junctions 


Gap junction 


Adjacent 
plasma 
membranes 


Gap between 
cells 


Connexons 
(composed of 
connexins) 


Adjacent 
plasma 

membranes 
Plaque 


Transmembrane 


Integrins 


Basal 
lamina ) ( 
Hemidesmosome 


glycoprotein glycoprotein 
(cadherin) (cadherin) 
Intermediate Actin filament 
filament 
(keratin) 
Intercellullar ——_-i Intercellullar 
space space _ 
it ee 
il 
Desmosome ( Adherens ) C) 
—— 


Adjacent 
plasma 
membranes 


Plaque 


Transmembrane 


The three basic types of cell-to-cell junctions are tight 
junctions, gap junctions, and anchoring junctions. 


At one end of the spectrum is the tight junction, which separates the cells 
into apical and basal compartments. When two adjacent epithelial cells 
form a tight junction, there is no extracellular space between them and the 
movement of substances through the extracellular space between the cells is 
blocked. This enables the epithelia to act as selective barriers. An 
anchoring junction includes several types of cell junctions that help 


stabilize epithelial tissues. Anchoring junctions are common on the lateral 
and basal surfaces of cells where they provide strong and flexible 
connections. There are three types of anchoring junctions: desmosomes, 
hemidesmosomes, and adherens. Desmosomes occur in patches on the 
membranes of cells. The patches are structural proteins on the inner surface 
of the cell’s membrane. The adhesion molecule, cadherin, is embedded in 
these patches and projects through the cell membrane to link with the 
cadherin molecules of adjacent cells. These connections are especially 
important in holding cells together. Hemidesmosomes, which look like half 
a desmosome, link cells to the extracellular matrix, for example, the basal 
lamina. While similar in appearance to desmosomes, they include the 
adhesion proteins called integrins rather than cadherins. Adherens junctions 
use either cadherins or integrins depending on whether they are linking to 
other cells or matrix. The junctions are characterized by the presence of the 
contractile protein actin located on the cytoplasmic surface of the cell 
membrane. The actin can connect isolated patches or form a belt-like 
structure inside the cell. These junctions influence the shape and folding of 
the epithelial tissue. 


In contrast with the tight and anchoring junctions, a gap junction forms an 
intercellular passageway between the membranes of adjacent cells to 
facilitate the movement of small molecules and ions between the cytoplasm 
of adjacent cells. These junctions allow electrical and metabolic coupling of 
adjacent cells, which coordinates function in large groups of cells. 


Classification of Epithelial Tissues 


Epithelial tissues are classified according to the shape of the cells and 
number of the cell layers formed ((link]). Cell shapes can be squamous 
(flattened and thin), cuboidal (boxy, as wide as it is tall), or columnar 
(rectangular, taller than it is wide). Similarly, the number of cell layers in 
the tissue can be one—where every cell rests on the basal lamina—which is 
a simple epithelium, or more than one, which is a stratified epithelium and 
only the basal layer of cells rests on the basal lamina. Pseudostratified 
(pseudo- = “false”) describes tissue with a single layer of irregularly shaped 
cells that give the appearance of more than one layer. Transitional describes 


a form of specialized stratified epithelium in which the shape of the cells 

can vary. 

Cells of Epithelial Tissue 
[Simp 


Stratified 


Squamous 


—S—S==—= 


Simple squamous epithelium 


Cuboidal elelelelele 


Simple cuboidal epithelium Stratified cuboidal epithelium Pseudostratified 


Simple columnar epithelium Stratified columnar epithelium Pseudostratified columnar epithelium 


Simple epithelial tissue is organized as a single layer of 
cells and stratified epithelial tissue is formed by several 
layers of cells. 


Simple Epithelium 


The shape of the cells in the single cell layer of simple epithelium reflects 
the functioning of those cells. The cells in simple squamous epithelium 
have the appearance of thin scales. Squamous cell nuclei tend to be flat, 
horizontal, and elliptical, mirroring the form of the cell. The endothelium 
is the epithelial tissue that lines vessels of the lymphatic and cardiovascular 


system, and it is made up of a single layer of squamous cells. Simple 
Squamous epithelium, because of the thinness of the cell, is present where 
rapid passage of chemical compounds is observed. The alveoli of lungs 
where gases diffuse, segments of kidney tubules, and the lining of 
capillaries are also made of simple squamous epithelial tissue. The 
mesothelium is a simple squamous epithelium that forms the surface layer 
of the serous membrane that lines body cavities and internal organs. Its 
primary function is to provide a smooth and protective surface. Mesothelial 
cells are squamous epithelial cells that secrete a fluid that lubricates the 
mesothelium. 


In simple cuboidal epithelium, the nucleus of the box-like cells appears 
round and is generally located near the center of the cell. These epithelia are 
active in the secretion and absorptions of molecules. Simple cuboidal 
epithelia are observed in the lining of the kidney tubules and in the ducts of 
glands. 


In simple columnar epithelium, the nucleus of the tall column-like cells 
tends to be elongated and located in the basal end of the cells. Like the 
cuboidal epithelia, this epithelium is active in the absorption and secretion 
of molecules. Simple columnar epithelium forms the lining of some 
sections of the digestive system and parts of the female reproductive tract. 
Ciliated columnar epithelium is composed of simple columnar epithelial 
cells with cilia on their apical surfaces. These epithelial cells are found in 
the lining of the fallopian tubes and parts of the respiratory system, where 
the beating of the cilia helps remove particulate matter. 


Pseudostratified columnar epithelium is a type of epithelium that appears 
to be stratified but instead consists of a single layer of irregularly shaped 
and differently sized columnar cells. In pseudostratified epithelium, nuclei 
of neighboring cells appear at different levels rather than clustered in the 
basal end. The arrangement gives the appearance of stratification; but in 
fact all the cells are in contact with the basal lamina, although some do not 
reach the apical surface. Pseudostratified columnar epithelium is found in 
the respiratory tract, where some of these cells have cilia. 


Both simple and pseudostratified columnar epithelia are heterogeneous 
epithelia because they include additional types of cells interspersed among 


the epithelial cells. For example, a goblet cell is a mucous-secreting 
unicellular “gland” interspersed between the columnar epithelial cells of 
mucous membranes ([link]). 

Goblet Cell 


Microvilli 


Secretory vesicles 
containing mucin 


Nucleus 


(a) In the lining of the small intestine, columnar 
epithelium cells are interspersed with goblet cells. (b) 
The arrows in this micrograph point to the mucous- 
secreting goblet cells. LM x 1600. (Micrograph 
provided by the Regents of University of Michigan 
Medical School © 2012) 


Note: 


CEs.) 
a 


— 
mss Openstax COLLEGE 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. 


Stratified Epithelium 


A stratified epithelium consists of several stacked layers of cells. This 
epithelium protects against physical and chemical wear and tear. The 
stratified epithelium is named by the shape of the most apical layer of cells, 
closest to the free space. Stratified squamous epithelium is the most 
common type of stratified epithelium in the human body. The apical cells 
are squamous, whereas the basal layer contains either columnar or cuboidal 
cells. The top layer may be covered with dead cells filled with keratin. 
Mammalian skin is an example of this dry, keratinized, stratified squamous 
epithelium. The lining of the mouth cavity is an example of an 
unkeratinized, stratified squamous epithelium. Stratified cuboidal 
epithelium and stratified columnar epithelium can also be found in 
certain glands and ducts, but are uncommon in the human body. 


Another kind of stratified epithelium is transitional epithelium, so-called 
because of the gradual changes in the shapes of the apical cells as the 
bladder fills with urine. It is found only in the urinary system, specifically 
the ureters and urinary bladder. When the bladder is empty, this epithelium 
is convoluted and has cuboidal apical cells with convex, umbrella shaped, 
apical surfaces. As the bladder fills with urine, this epithelium loses its 
convolutions and the apical cells transition from cuboidal to squamous. It 
appears thicker and more multi-layered when the bladder is empty, and 
more stretched out and less stratified when the bladder is full and distended. 
[link] summarizes the different categories of epithelial cell tissue cells. 
Summary of Epithelial Tissue Cells 


Simple cuboidal epithelium 


Stratified cuboidal epithelium 


Air sacs of lungs and the lining 
of the heart, blood vessels, 
and lymphatic vessels 


In ducts and secretory portions 
of small glands and in kidney 
tubules 


Ciliated tissues are in bronchi, 
uterine tubes, and uterus; 
smooth (nonciliated tissues) 
are in the digestive tract, 
bladder 


Ciliated tissue lines the trachea 
and much of the upper 
respiratory tract 


Lines the esophagus, mouth, 
and vagina 


Sweat glands, salivary glands, 
and the mammary glands 


The male urethra and the 


ducts of some glands 


Lines the bladder, uretha, and 
the ureters 


SS 


Simple squamous epithelium 


Allows materials to pass 
through by diffusion and 
filtration, and secretes 
lubricating substance 


Secretes and absorbs 


Absorbs; it also secretes 
mucous and enzymes 


Secretes mucus; ciliated tissue 
moves mucus 


Protects against abrasion 


Protective tissue 


Secretes and protects 


Allows the urinary organs to 
expand and stretch 


Watch this video to find out more about the anatomy of epithelial tissues. 
Where in the body would one find non-keratinizing stratified squamous 
epithelium? 


Glandular Epithelium 


A gland is a structure made up of one or more cells modified to synthesize 
and secrete chemical substances. Most glands consist of groups of epithelial 
cells. A gland can be classified as an endocrine gland, a ductless gland that 
releases secretions directly into surrounding tissues and fluids (endo- = 
“inside”), or an exocrine gland whose secretions leave through a duct that 
opens directly, or indirectly, to the external environment (exo- = “outside’”’). 


Endocrine Glands 


The secretions of endocrine glands are called hormones. Hormones are 
released into the interstitial fluid, diffused into the bloodstream, and 
delivered to targets, in other words, cells that have receptors to bind the 
hormones. The endocrine system is part of a major regulatory system 
coordinating the regulation and integration of body responses. A few 
examples of endocrine glands include the anterior pituitary, thymus, adrenal 
cortex, and gonads. 


Exocrine Glands 


Exocrine glands release their contents through a duct that leads to the 
epithelial surface. Mucous, sweat, saliva, and breast milk are all examples 
of secretions from exocrine glands. They are all discharged through tubular 
ducts. Secretions into the lumen of the gastrointestinal tract, technically 
outside of the body, are of the exocrine category. 


Glandular Structure 


Exocrine glands are classified as either unicellular or multicellular. The 
unicellular glands are scattered single cells, such as goblet cells, found in 
the mucous membranes of the small and large intestine. 


The multicellular exocrine glands known as serous glands develop from 
simple epithelium to form a secretory surface that secretes directly into an 
inner cavity. These glands line the internal cavities of the abdomen and 
chest and release their secretions directly into the cavities. Other 
multicellular exocrine glands release their contents through a tubular duct. 
The duct is single in a simple gland but in compound glands is divided into 
one or more branches ([link]). In tubular glands, the ducts can be straight or 
coiled, whereas tubes that form pockets are alveolar (acinar), such as the 
exocrine portion of the pancreas. Combinations of tubes and pockets are 
known as tubuloalveolar (tubuloacinar) compound glands. In a branched 
gland, a duct is connected to more than one secretory group of cells. 

Types of Exocrine Glands 


Simple ducts 


a 


Alveolar 
(acinar) 


Not found in 
adult; a stage in 
development of 
simple 
branched 
glands 


Simp! 


Sebaceous (oil) 
glands 


Compound alveolar (acinar) 


RRRROESSAS 


Compound tubuloalveolar 


> 
o 
ix y 
ing 
a 


- 


Salivary glands; glands of 
respiratory passages; 
and pancreas 


Mammary glands 


Simple tubular — 


Simple branched tubular 
aTIr- - AP 


Intestinal glands 


Merocrine sweat 
glands 


Gastric glands; 

and mucous glands 
of esophagus, tongue, 
duodenum 


Compound tubular 


DOOUDWODLE ot ee oes 


Mucous glands (in mouth); 
bulbourethral glands (in male 
reproductive system); and testes 


(seminiferous tubules) ) 
Compound ducts 4 


Exocrine glands are classified by their structure. 


Methods and Types of Secretion 


Exocrine glands can be classified by their mode of secretion and the nature 


Tubular 


of the substances released, as well as by the structure of the glands and 


shape of ducts ([link]). Merocrine secretion is the most common type of 
exocrine secretion. The secretions are enclosed in vesicles that move to the 
apical surface of the cell where the contents are released by exocytosis. For 
example, watery mucous containing the glycoprotein mucin, a lubricant that 
offers some pathogen protection is a merocrine secretion. The eccrine 
glands that produce and secrete sweat are another example. 

Modes of Glandular Secretion 


Secretion 


Secretory vesicle 


(a) Merocrine Golgi complex 


secretion 


Nucleus 


Pinched off portion 
of cell is the secretion 


(b) Apocrine 
secretion 


Mature cell dies 
and becomes 
secretory product 


(c) Holocrine 
secretion 


(a) In merocrine secretion, the cell remains 
intact. (b) In apocrine secretion, the apical 
portion of the cell is released, as well. (c) In 
holocrine secretion, the cell is destroyed as it 
releases its product and the cell itself 
becomes part of the secretion. 


Apocrine secretion accumulates near the apical portion of the cell. That 
portion of the cell and its secretory contents pinch off from the cell and are 


released. Apocrine sweat glands in the axillary and genital areas release 
fatty secretions that local bacteria break down; this causes body odor. Both 
merocrine and apocrine glands continue to produce and secrete their 
contents with little damage caused to the cell because the nucleus and golgi 
regions remain intact after secretion. 


In contrast, the process of holocrine secretion involves the rupture and 
destruction of the entire gland cell. The cell accumulates its secretory 
products and releases them only when it bursts. New gland cells 
differentiate from cells in the surrounding tissue to replace those lost by 
secretion. The sebaceous glands that produce the oils on the skin and hair 
are holocrine glands/cells ({link]). 

Sebaceous Glands 


Hair 


Sebaceous 
gland 


These glands secrete oils that lubricate and protect the skin. 
They are holocrine glands and they are destroyed after 
releasing their contents. New glandular cells form to 
replace the cells that are lost. LM x 400. (Micrograph 
provided by the Regents of University of Michigan Medical 
School © 2012) 


Glands are also named after the products they produce. The serous gland 
produces watery, blood-plasma-like secretions rich in enzymes such as 


alpha amylase, whereas the mucous gland releases watery to viscous 
products rich in the glycoprotein mucin. Both serous and mucous glands are 
common in the salivary glands of the mouth. Mixed exocrine glands contain 
both serous and mucous glands and release both types of secretions. 


Chapter Review 


In epithelial tissue, cells are closely packed with little or no extracellular 
matrix except for the basal lamina that separates the epithelium from 
underlying tissue. The main functions of epithelia are protection from the 
environment, coverage, secretion and excretion, absorption, and filtration. 
Cells are bound together by tight junctions that form an impermeable 
barrier. They can also be connected by gap junctions, which allow free 
exchange of soluble molecules between cells, and anchoring junctions, 
which attach cell to cell or cell to matrix. The different types of epithelial 
tissues are characterized by their cellular shapes and arrangements: 
squamous, cuboidal, or columnar epithelia. Single cell layers form simple 
epithelia, whereas stacked cells form stratified epithelia. Very few 
capillaries penetrate these tissues. 


Glands are secretory tissues and organs that are derived from epithelial 
tissues. Exocrine glands release their products through ducts. Endocrine 
glands secrete hormones directly into the interstitial fluid and blood stream. 
Glands are classified both according to the type of secretion and by their 
structure. Merocrine glands secrete products as they are synthesized. 
Apocrine glands release secretions by pinching off the apical portion of the 
cell, whereas holocrine gland cells store their secretions until they rupture 
and release their contents. In this case, the cell becomes part of the 
secretion. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to find out more about the anatomy of epithelial 
tissues. Where in the body would one find non-keratinizing stratified 
squamous epithelium? 


Solution: 


The inside of the mouth, esophagus, vaginal canal, and anus. 


Review Questions 


Exercise: 
Problem: 
In observing epithelial cells under a microscope, the cells are arranged 
in a single layer and look tall and narrow, and the nucleus is located 


close to the basal side of the cell. The specimen is what type of 
epithelial tissue? 


a. columnar 
b. stratified 

c. squamous 
d. transitional 


Solution: 


A 
Exercise: 


Problem: 


Which of the following is the epithelial tissue that lines the interior of 
blood vessels? 


a. columnar 


b. pseudostratified 
c. simple squamous 
d. transitional 


Solution: 


C 
Exercise: 
Problem: 


Which type of epithelial tissue specializes in moving particles across 
its surface and is found in airways and lining of the oviduct? 


a. transitional 

b. stratified columnar 

c. pseudostratified ciliated columnar 
d. stratified squamous 


Solution: 


B 
Exercise: 
Problem: 
The exocrine gland stores its secretion until the glandular 


cell ruptures, whereas the gland releases its apical region 
and reforms. 


a. holocrine; apocrine 
b. eccrine; endocrine 
c. apocrine; holocrine 
d. eccrine; apocrine 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: 


The structure of a tissue usually is optimized for its function. Describe 
how the structure of the mucosa and its cells match its function of 
nutrient absorption. 


Solution: 


The mucosa of the intestine is highly folded, increasing the surface 
area for nutrient absorption. A greater surface area for absorption 
allows more nutrients to be absorbed per unit time. In addition, the 
nutrient-absorbing cells of the mucosa have finger-like projections 
called microvilli that further increase the surface area for nutrient 
absorption. 


Glossary 


anchoring junction 
mechanically attaches adjacent cells to each other or to the basement 
membrane 


apical 
that part of a cell or tissue which, in general, faces an open space 


apocrine secretion 
release of a substance along with the apical portion of the cell 


basal lamina 
thin extracellular layer that lies underneath epithelial cells and 
separates them from other tissues 


basement membrane 
in epithelial tissue, a thin layer of fibrous material that anchors the 
epithelial tissue to the underlying connective tissue; made up of the 
basal lamina and reticular lamina 


cell junction 
point of cell-to-cell contact that connects one cell to another in a tissue 


endocrine gland 
groups of cells that release chemical signals into the intercellular fluid 
to be picked up and transported to their target organs by blood 


endothelium 
tissue that lines vessels of the lymphatic and cardiovascular system, 
made up of a simple squamous epithelium 


exocrine gland 
group of epithelial cells that secrete substances through ducts that open 
to the skin or to internal body surfaces that lead to the exterior of the 
body 


gap junction 
allows cytoplasmic communications to occur between cells 


goblet cell 
unicellular gland found in columnar epithelium that secretes mucous 


holocrine secretion 
release of a substance caused by the rupture of a gland cell, which 
becomes part of the secretion 


merocrine secretion 
release of a substance from a gland via exocytosis 


mesothelium 
simple squamous epithelial tissue which covers the major body 
cavities and is the epithelial portion of serous membranes 


mucous gland 
group of cells that secrete mucous, a thick, slippery substance that 
keeps tissues moist and acts as a lubricant 


pseudostratified columnar epithelium 
tissue that consists of a single layer of irregularly shaped and sized 
cells that give the appearance of multiple layers; found in ducts of 
certain glands and the upper respiratory tract 


reticular lamina 
matrix containing collagen and elastin secreted by connective tissue; a 
component of the basement membrane 


serous gland 
group of cells within the serous membrane that secrete a lubricating 
substance onto the surface 


simple columnar epithelium 
tissue that consists of a single layer of column-like cells; promotes 
secretion and absorption in tissues and organs 


simple cuboidal epithelium 
tissue that consists of a single layer of cube-shaped cells; promotes 
secretion and absorption in ducts and tubules 


simple squamous epithelium 
tissue that consists of a single layer of flat scale-like cells; promotes 
diffusion and filtration across surface 


stratified columnar epithelium 
tissue that consists of two or more layers of column-like cells, contains 
glands and is found in some ducts 


stratified cuboidal epithelium 
tissue that consists of two or more layers of cube-shaped cells, found 


in some ducts 


stratified squamous epithelium 


tissue that consists of multiple layers of cells with the most apical 
being flat scale-like cells; protects surfaces from abrasion 


tight junction 
forms an impermeable barrier between cells 


transitional epithelium 
form of stratified epithelium found in the urinary tract, characterized 
by an apical layer of cells that change shape in response to the 
presence of urine 


Connective Tissue Supports and Protects 
By the end of this section, you will be able to: 


e Identify and distinguish between the types of connective tissue: proper, 
supportive, and fluid 
e Explain the functions of connective tissues 


As may be obvious from its name, one of the major functions of connective 
tissue is to connect tissues and organs. Unlike epithelial tissue, which is 
composed of cells closely packed with little or no extracellular space in 
between, connective tissue cells are dispersed in a matrix. The matrix 
usually includes a large amount of extracellular material produced by the 
connective tissue cells that are embedded within it. The matrix plays a 
major role in the functioning of this tissue. The major component of the 
matrix is a ground substance often crisscrossed by protein fibers. This 
ground substance is usually a fluid, but it can also be mineralized and solid, 
as in bones. Connective tissues come in a vast variety of forms, yet they 
typically have in common three characteristic components: cells, large 
amounts of amorphous ground substance, and protein fibers. The amount 
and structure of each component correlates with the function of the tissue, 
from the rigid ground substance in bones supporting the body to the 
inclusion of specialized cells; for example, a phagocytic cell that engulfs 
pathogens and also rids tissue of cellular debris. 


Functions of Connective Tissues 


Connective tissues perform many functions in the body, but most 
importantly, they support and connect other tissues; from the connective 
tissue sheath that surrounds muscle cells, to the tendons that attach muscles 
to bones, and to the skeleton that supports the positions of the body. 
Protection is another major function of connective tissue, in the form of 
fibrous capsules and bones that protect delicate organs and, of course, the 
skeletal system. Specialized cells in connective tissue defend the body from 
microorganisms that enter the body. Transport of fluid, nutrients, waste, and 
chemical messengers is ensured by specialized fluid connective tissues, 
such as blood and lymph. Adipose cells store surplus energy in the form of 
fat and contribute to the thermal insulation of the body. 


Embryonic Connective Tissue 


All connective tissues derive from the mesodermal layer of the embryo (see 
[link]). The first connective tissue to develop in the embryo is 
mesenchyme, the stem cell line from which all connective tissues are later 
derived. Clusters of mesenchymal cells are scattered throughout adult tissue 
and supply the cells needed for replacement and repair after a connective 
tissue injury. A second type of embryonic connective tissue forms in the 
umbilical cord, called mucous connective tissue or Wharton’s jelly. This 
tissue is no longer present after birth, leaving only scattered mesenchymal 
cells throughout the body. 


Classification of Connective Tissues 


The three broad categories of connective tissue are classified according to 
the characteristics of their ground substance and the types of fibers found 
within the matrix ({link]). Connective tissue proper includes loose 
connective tissue and dense connective tissue. Both tissues have a variety 
of cell types and protein fibers suspended in a viscous ground substance. 
Dense connective tissue is reinforced by bundles of fibers that provide 
tensile strength, elasticity, and protection. In loose connective tissue, the 
fibers are loosely organized, leaving large spaces in between. Supportive 
connective tissue—bone and cartilage—provide structure and strength to 
the body and protect soft tissues. A few distinct cell types and densely 
packed fibers in a matrix characterize these tissues. In bone, the matrix is 
rigid and described as calcified because of the deposited calcium salts. In 
fluid connective tissue, in other words, lymph and blood, various 
specialized cells circulate in a watery fluid containing salts, nutrients, and 
dissolved proteins. 


Connective Tissue Examples 


Connective tisssiee Exaifipjasortive Fluid connective 


proper connective tissue tissue 
Connective tissue Supportive Fluid connective 
proper connective tissue tissue 
Loose connective : 
an Cartilage 
e Hyaline 
e Areolar y Blood 
a aadinose e Fibrocartilage 
e Reticular Pane 
Dense connective 
tissue 
Bones 
Regular 
; ere e Compact bone Lymph 
eicsalat e Cancellous bone 
elastic 


Connective Tissue Proper 


Fibroblasts are present in all connective tissue proper ([link]). Fibrocytes, 
adipocytes, and mesenchymal cells are fixed cells, which means they 
remain within the connective tissue. Other cells move in and out of the 
connective tissue in response to chemical signals. Macrophages, mast cells, 
lymphocytes, plasma cells, and phagocytic cells are found in connective 
tissue proper but are actually part of the immune system protecting the 
body. 

Connective Tissue Proper 


Reticular fibers 
Adipocytes 
Mesenchymal cell 
Elastic fibers 
Collagen fibers 
Fibroblast 


Macrophage 


Fibroblasts produce this fibrous tissue. Connective tissue proper 
includes the fixed cells fibrocytes, adipocytes, and mesenchymal cells. 
LM x 400. (Micrograph provided by the Regents of University of 
Michigan Medical School © 2012) 


Cell Types 


The most abundant cell in connective tissue proper is the fibroblast. 
Polysaccharides and proteins secreted by fibroblasts combine with extra- 
cellular fluids to produce a viscous ground substance that, with embedded 
fibrous proteins, forms the extra-cellular matrix. As you might expect, a 
fibrocyte, a less active form of fibroblast, is the second most common cell 
type in connective tissue proper. 


Adipocytes are cells that store lipids as droplets that fill most of the 
cytoplasm. There are two basic types of adipocytes: white and brown. The 
brown adipocytes store lipids as many droplets, and have high metabolic 
activity. In contrast, white fat adipocytes store lipids as a single large drop 
and are metabolically less active. Their effectiveness at storing large 
amounts of fat is witnessed in obese individuals. The number and type of 
adipocytes depends on the tissue and location, and vary among individuals 
in the population. 


The mesenchymal cell is a multipotent adult stem cell. These cells can 
differentiate into any type of connective tissue cells needed for repair and 
healing of damaged tissue. 


The macrophage cell is a large cell derived from a monocyte, a type of 
blood cell, which enters the connective tissue matrix from the blood vessels. 
The macrophage cells are an essential component of the immune system, 
which is the body’s defense against potential pathogens and degraded host 
cells. When stimulated, macrophages release cytokines, small proteins that 
act as chemical messengers. Cytokines recruit other cells of the immune 
system to infected sites and stimulate their activities. Roaming, or free, 
macrophages move rapidly by amoeboid movement, engulfing infectious 
agents and cellular debris. In contrast, fixed macrophages are permanent 
residents of their tissues. 


The mast cell, found in connective tissue proper, has many cytoplasmic 
granules. These granules contain the chemical signals histamine and 
heparin. When irritated or damaged, mast cells release histamine, an 
inflammatory mediator, which causes vasodilation and increased blood flow 
at a site of injury or infection, along with itching, swelling, and redness you 
recognize as an allergic response. Like blood cells, mast cells are derived 
from hematopoietic stem cells and are part of the immune system. 


Connective Tissue Fibers and Ground Substance 


Three main types of fibers are secreted by fibroblasts: collagen fibers, 
elastic fibers, and reticular fibers. Collagen fiber is made from fibrous 
protein subunits linked together to form a long and straight fiber. Collagen 
fibers, while flexible, have great tensile strength, resist stretching, and give 
ligaments and tendons their characteristic resilience and strength. These 
fibers hold connective tissues together, even during the movement of the 
body. 


Elastic fiber contains the protein elastin along with lesser amounts of other 
proteins and glycoproteins. The main property of elastin is that after being 
stretched or compressed, it will return to its original shape. Elastic fibers are 


prominent in elastic tissues found in skin and the elastic ligaments of the 
vertebral column. 


Reticular fiber is also formed from the same protein subunits as collagen 
fibers; however, these fibers remain narrow and are arrayed in a branching 
network. They are found throughout the body, but are most abundant in the 
reticular tissue of soft organs, such as liver and spleen, where they anchor 
and provide structural support to the parenchyma (the functional cells, 
blood vessels, and nerves of the organ). 


All of these fiber types are embedded in ground substance. Secreted by 
fibroblasts, ground substance is made of polysaccharides, specifically 
hyaluronic acid, and proteins. These combine to form a proteoglycan with a 
protein core and polysaccharide branches. The proteoglycan attracts and 
traps available moisture forming the clear, viscous, colorless matrix you 
now know as ground substance. 


Loose Connective Tissue 


Loose connective tissue is found between many organs where it acts both to 
absorb shock and bind tissues together. It allows water, salts, and various 
nutrients to diffuse through to adjacent or imbedded cells and tissues. 


Adipose tissue consists mostly of fat storage cells, with little extracellular 
matrix ({link]). A large number of capillaries allow rapid storage and 
mobilization of lipid molecules. White adipose tissue is most abundant. It 
can appear yellow and owes its color to carotene and related pigments from 
plant food. White fat contributes mostly to lipid storage and can serve as 
insulation from cold temperatures and mechanical injuries. White adipose 
tissue can be found protecting the kidneys and cushioning the back of the 
eye. Brown adipose tissue is more common in infants, hence the term “baby 
fat.” In adults, there is a reduced amount of brown fat and it is found mainly 
in the neck and clavicular regions of the body. The many mitochondria in 
the cytoplasm of brown adipose tissue help explain its efficiency at 
metabolizing stored fat. Brown adipose tissue is thermogenic, meaning that 


as it breaks down fats, it releases metabolic heat, rather than producing 
adenosine triphosphate (ATP), a key molecule used in metabolism. 
Adipose Tissue 


| es ————— 


This is a loose connective tissue that consists of fat cells 
with little extracellular matrix. It stores fat for energy 
and provides insulation. LM x 800. (Micrograph 
provided by the Regents of University of Michigan 
Medical School © 2012) 


Areolar tissue shows little specialization. It contains all the cell types and 
fibers previously described and is distributed in a random, web-like fashion. 
It fills the spaces between muscle fibers, surrounds blood and lymph 
vessels, and supports organs in the abdominal cavity. Areolar tissue 
underlies most epithelia and represents the connective tissue component of 
epithelial membranes, which are described further in a later section. 


Reticular tissue is a mesh-like, supportive framework for soft organs such 
as lymphatic tissue, the spleen, and the liver ([link]). Reticular cells produce 
the reticular fibers that form the network onto which other cells attach. It 
derives its name from the Latin reticulus, which means “little net.” 
Reticular Tissue 


This is a loose connective tissue made up of a network of 
reticular fibers that provides a supportive framework for 
soft organs. LM x 1600. (Micrograph provided by the 
Regents of University of Michigan Medical School © 
2012) 


Dense Connective Tissue 


Dense connective tissue contains more collagen fibers than does loose 
connective tissue. As a consequence, it displays greater resistance to 
stretching. There are two major categories of dense connective tissue: 
regular and irregular. Dense regular connective tissue fibers are parallel to 
each other, enhancing tensile strength and resistance to stretching in the 
direction of the fiber orientations. Ligaments and tendons are made of dense 
regular connective tissue, but in ligaments not all fibers are parallel. Dense 
regular elastic tissue contains elastin fibers in addition to collagen fibers, 
which allows the ligament to return to its original length after stretching. 
The ligaments in the vocal folds and between the vertebrae in the vertebral 
column are elastic. 


In dense irregular connective tissue, the direction of fibers is random. This 
arrangement gives the tissue greater strength in all directions and less 
strength in one particular direction. In some tissues, fibers crisscross and 
form a mesh. In other tissues, stretching in several directions is achieved by 


alternating layers where fibers run in the same orientation in each layer, and 
it is the layers themselves that are stacked at an angle. The dermis of the 
skin is an example of dense irregular connective tissue rich in collagen 
fibers. Dense irregular elastic tissues give arterial walls the strength and the 
ability to regain original shape after stretching ([link]). 

Dense Connective Tissue 


Fibroblast 
nuclei 


Fibroblast 
nuclei 


Collagen 
fiber 
bundles 


(b) Irregular dense 


(a) Dense regular connective tissue consists of 
collagenous fibers packed into parallel bundles. (b) 
Dense irregular connective tissue consists of collagenous 
fibers interwoven into a mesh-like network. From top, 
LM x 1000, LM x 200. (Micrographs provided by the 
Regents of University of Michigan Medical School © 
2012) 


Note: 
Disorders of the... 
Connective Tissue: Tendinitis 


Your opponent stands ready as you prepare to hit the serve, but you are 
confident that you will smash the ball past your opponent. As you toss the 
ball high in the air, a bumming pain shoots across your wrist and you drop 
the tennis racket. That dull ache in the wrist that you ignored through the 
summer is now an unbearable pain. The game is over for now. 

After examining your swollen wrist, the doctor in the emergency room 
announces that you have developed wrist tendinitis. She recommends icing 
the tender area, taking non-steroidal anti-inflammatory medication to ease 
the pain and to reduce swelling, and complete rest for a few weeks. She 
interrupts your protests that you cannot stop playing. She issues a stern 
warning about the risk of aggravating the condition and the possibility of 
surgery. She consoles you by mentioning that well known tennis players 
such as Venus and Serena Williams and Rafael Nadal have also suffered 
from tendinitis related injuries. 

What is tendinitis and how did it happen? Tendinitis is the inflammation of 
a tendon, the thick band of fibrous connective tissue that attaches a muscle 
to a bone. The condition causes pain and tenderness in the area around a 
joint. On rare occasions, a sudden serious injury will cause tendinitis. Most 
often, the condition results from repetitive motions over time that strain the 
tendons needed to perform the tasks. 

Persons whose jobs and hobbies involve performing the same movements 
over and over again are often at the greatest risk of tendinitis. You hear of 
tennis and golfer’s elbow, jumper's knee, and swimmer’s shoulder. In all 
cases, overuse of the joint causes a microtrauma that initiates the 
inflammatory response. Tendinitis is routinely diagnosed through a clinical 
examination. In case of severe pain, X-rays can be examined to rule out the 
possibility of a bone injury. Severe cases of tendinitis can even tear loose a 
tendon. Surgical repair of a tendon is painful. Connective tissue in the 
tendon does not have abundant blood supply and heals slowly. 

While older adults are at risk for tendinitis because the elasticity of tendon 
tissue decreases with age, active people of all ages can develop tendinitis. 
Young athletes, dancers, and computer operators; anyone who performs the 
same movements constantly is at risk for tendinitis. Although repetitive 
motions are unavoidable in many activities and may lead to tendinitis, 
precautions can be taken that can lessen the probability of developing 
tendinitis. For active individuals, stretches before exercising and cross 
training or changing exercises are recommended. For the passionate 


athlete, it may be time to take some lessons to improve technique. All of 
the preventive measures aim to increase the strength of the tendon and 
decrease the stress put on it. With proper rest and managed care, you will 
be back on the court to hit that slice-spin serve over the net. 


Note: 


Pelee Le 


Watch this animation to learn more about tendonitis, a painful condition 
caused by swollen or injured tendons. 


Supportive Connective Tissues 


Two major forms of supportive connective tissue, cartilage and bone, allow 
the body to maintain its posture and protect internal organs. 


Cartilage 


The distinctive appearance of cartilage is due to polysaccharides called 
chondroitin sulfates, which bind with ground substance proteins to form 
proteoglycans. Embedded within the cartilage matrix are chondrocytes, or 
cartilage cells, and the space they occupy are called lacunae (singular = 
lacuna). A layer of dense irregular connective tissue, the perichondrium, 
encapsulates the cartilage. Cartilaginous tissue is avascular, thus all 
nutrients need to diffuse through the matrix to reach the chondrocytes. This 
is a factor contributing to the very slow healing of cartilaginous tissues. 


The three main types of cartilage tissue are hyaline cartilage, fibrocartilage, 
and elastic cartilage ([{link]). Hyaline cartilage, the most common type of 
cartilage in the body, consists of short and dispersed collagen fibers and 
contains large amounts of proteoglycans. Under the microscope, tissue 
samples appear clear. The surface of hyaline cartilage is smooth. Both 
strong and flexible, it is found in the rib cage and nose and covers bones 
where they meet to form moveable joints. It makes up a template of the 
embryonic skeleton before bone formation. A plate of hyaline cartilage at 
the ends of bone allows continued growth until adulthood. Fibrocartilage 
is tough because it has thick bundles of collagen fibers dispersed through its 
matrix. Menisci in the knee joint and the intervertebral discs are examples 
of fibrocartilage. Elastic cartilage contains elastic fibers as well as collagen 
and proteoglycans. This tissue gives rigid support as well as elasticity. Tug 
gently at your ear lobes, and notice that the lobes return to their initial 
shape. The external ear contains elastic cartilage. 

Types of Cartilage 


(a) Hyaline cartilage 


Chondrocytes 
in lacunae 


Matrix 


(b) Fibrocartilage 


Chondrocyte 
in lacuna 


Collagen fiber 
in matrix 


(c) Elastic cartilage 


Chondrocyte 
in lacuna 


Elastic fibers 
in matrix 


Cartilage is a connective tissue consisting of 
collagenous fibers embedded in a firm matrix of 
chondroitin sulfates. (a) Hyaline cartilage provides 
support with some flexibility. The example is from 
dog tissue. (b) Fibrocartilage provides some 
compressibility and can absorb pressure. (c) Elastic 
cartilage provides firm but elastic support. From top, 
LM x 300, LM x 1200, LM x 1016. (Micrographs 
provided by the Regents of University of Michigan 
Medical School © 2012) 


Bone 


Bone is the hardest connective tissue. It provides protection to internal 
organs and supports the body. Bone’s rigid extracellular matrix contains 
mostly collagen fibers embedded in a mineralized ground substance 
containing hydroxyapatite, a form of calcium phosphate. Both components 
of the matrix, organic and inorganic, contribute to the unusual properties of 
bone. Without collagen, bones would be brittle and shatter easily. Without 
mineral crystals, bones would flex and provide little support. Osteocytes, 
bone cells like chondrocytes, are located within lacunae. The histology of 
transverse tissue from long bone shows a typical arrangement of osteocytes 
in concentric circles around a central canal. Bone is a highly vascularized 
tissue. Unlike cartilage, bone tissue can recover from injuries in a relatively 
short time. 


Cancellous bone looks like a sponge under the microscope and contains 
empty spaces between trabeculae, or arches of bone proper. It is lighter than 
compact bone and found in the interior of some bones and at the end of long 
bones. Compact bone is solid and has greater structural strength. 


Fluid Connective Tissue 


Blood and lymph are fluid connective tissues. Cells circulate in a liquid 
extracellular matrix. The formed elements circulating in blood are all 
derived from hematopoietic stem cells located in bone marrow ([link]). 
Erythrocytes, red blood cells, transport oxygen and some carbon dioxide. 
Leukocytes, white blood cells, are responsible for defending against 
potentially harmful microorganisms or molecules. Platelets are cell 
fragments involved in blood clotting. Some white blood cells have the 
ability to cross the endothelial layer that lines blood vessels and enter 
adjacent tissues. Nutrients, salts, and wastes are dissolved in the liquid 
matrix and transported through the body. 


Lymph contains a liquid matrix and white blood cells. Lymphatic capillaries 
are extremely permeable, allowing larger molecules and excess fluid from 
interstitial spaces to enter the lymphatic vessels. Lymph drains into blood 
vessels, delivering molecules to the blood that could not otherwise directly 
enter the bloodstream. In this way, specialized lymphatic capillaries 
transport absorbed fats away from the intestine and deliver these molecules 
to the blood. 

Blood: A Fluid Connective Tissue 


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Blood is a fluid connective tissue containing 
erythrocytes and various types of leukocytes 
that circulate in a liquid extracellular matrix. 
LM x 1600. (Micrograph provided by the 
Regents of University of Michigan Medical 
School © 2012) 


Note: 


ac| 


openstax COLLEGE” 


Wii 


ae 


View the University of Michigan Webscope to explore the tissue sample in 
greater detail. 


Note: 


=" 
meee OFENStAX COLLEGE” 


Visit this link to test your connective tissue knowledge with this 10- 
question quiz. Can you name the 10 tissue types shown in the histology 
slides? 


Chapter Review 


Connective tissue is a heterogeneous tissue with many cell shapes and 
tissue architecture. Structurally, all connective tissues contain cells that are 
embedded in an extracellular matrix stabilized by proteins. The chemical 
nature and physical layout of the extracellular matrix and proteins vary 
enormously among tissues, reflecting the variety of functions that 
connective tissue fulfills in the body. Connective tissues separate and 
cushion organs, protecting them from shifting or traumatic injury. Connect 
tissues provide support and assist movement, store and transport energy 
molecules, protect against infections, and contribute to temperature 
homeostasis. 


Many different cells contribute to the formation of connective tissues. They 
originate in the mesodermal germ layer and differentiate from mesenchyme 
and hematopoietic tissue in the bone marrow. Fibroblasts are the most 
abundant and secrete many protein fibers, adipocytes specialize in fat 
storage, hematopoietic cells from the bone marrow give rise to all the blood 
cells, chondrocytes form cartilage, and osteocytes form bone. The 


extracellular matrix contains fluid, proteins, polysaccharide derivatives, 
and, in the case of bone, mineral crystals. Protein fibers fall into three major 
groups: collagen fibers that are thick, strong, flexible, and resist stretch; 
reticular fibers that are thin and form a supportive mesh; and elastin fibers 
that are thin and elastic. 


The major types of connective tissue are connective tissue proper, 
supportive tissue, and fluid tissue. Loose connective tissue proper includes 
adipose tissue, areolar tissue, and reticular tissue. These serve to hold 
organs and other tissues in place and, in the case of adipose tissue, isolate 
and store energy reserves. The matrix is the most abundant feature for loose 
tissue although adipose tissue does not have much extracellular matrix. 
Dense connective tissue proper is richer in fibers and may be regular, with 
fibers oriented in parallel as in ligaments and tendons, or irregular, with 
fibers oriented in several directions. Organ capsules (collagenous type) and 
walls of arteries (elastic type) contain dense irregular connective tissue. 
Cartilage and bone are supportive tissue. Cartilage contains chondrocytes 
and is somewhat flexible. Hyaline cartilage is smooth and clear, covers 
joints, and is found in the growing portion of bones. Fibrocartilage is tough 
because of extra collagen fibers and forms, among other things, the 
intervertebral discs. Elastic cartilage can stretch and recoil to its original 
shape because of its high content of elastic fibers. The matrix contains very 
few blood vessels. Bones are made of a rigid, mineralized matrix containing 
calcium salts, crystals, and osteocytes lodged in lacunae. Bone tissue is 
highly vascularized. Cancellous bone is spongy and less solid than compact 
bone. Fluid tissue, for example blood and lymph, is characterized by a 
liquid matrix and no supporting fibers. 


Interactive Link Questions 


Exercise: 
Problem: 
Visit this link to test your connective tissue knowledge with this 10- 


question quiz. Can you name the 10 tissue types shown in the 
histology slides? 


Solution: 


Click at the bottom of the quiz for the answers. 


Review Questions 


Exercise: 


Problem: 
Connective tissue is made of which three essential components? 


a. cells, ground substance, and carbohydrate fibers 
b. cells, ground substance, and protein fibers 

c. collagen, ground substance, and protein fibers 
d. matrix, ground substance, and fluid 


Solution: 


B 
Exercise: 


Problem: 


Under the microscope, a tissue specimen shows cells located in spaces 
scattered in a transparent background. This is probably 


a. loose connective tissue 
b. a tendon 

c. bone 

d. hyaline cartilage 


Solution: 


D 


Exercise: 


Problem: Which connective tissue specializes in storage of fat? 


a. tendon 

b. adipose tissue 

c. reticular tissue 

d. dense connective tissue 


Solution: 


B 
Exercise: 
Problem: 


Ligaments connect bones together and withstand a lot of stress. What 
type of connective tissue should you expect ligaments to contain? 


a. areolar tissue 

b. adipose tissue 

c. dense regular connective tissue 
d. dense irregular connective tissue 


Solution: 


C 
Exercise: 


Problem: 
In adults, new connective tissue cells originate from the 


a. mesoderm 
b. mesenchyme 
c. ectoderm 


d. endoderm 


Solution: 


B 


Exercise: 


Problem:In bone, the main cells are 


a. fibroblasts 
b. chondrocytes 
c. lymphocytes 
d. osteocytes 


Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: 


One of the main functions of connective tissue is to integrate organs 
and organ systems in the body. Discuss how blood fulfills this role. 


Solution: 


Blood is a fluid connective tissue, a variety of specialized cells that 
circulate in a watery fluid containing salts, nutrients, and dissolved 
proteins in a liquid extracellular matrix. Blood contains formed 
elements derived from bone marrow. Erythrocytes, or red blood cells, 
transport the gases oxygen and carbon dioxide. Leukocytes, or white 
blood cells, are responsible for the defense of the organism against 


potentially harmful microorganisms or molecules. Platelets are cell 
fragments involved in blood clotting. Some cells have the ability to 
cross the endothelial layer that lines vessels and enter adjacent tissues. 
Nutrients, salts, and waste are dissolved in the liquid matrix and 
transported through the body. 


Exercise: 
Problem: 


Why does an injury to cartilage, especially hyaline cartilage, heal 
much more slowly than a bone fracture? 


Solution: 


A layer of dense irregular connective tissue covers cartilage. No blood 
vessels supply cartilage tissue. Injuries to cartilage heal very slowly 
because cells and nutrients needed for repair diffuse slowly to the 
injury site. 


Glossary 


adipocytes 
lipid storage cells 


adipose tissue 
specialized areolar tissue rich in stored fat 


areolar tissue 
(also, loose connective tissue) a type of connective tissue proper that 
shows little specialization with cells dispersed in the matrix 


chondrocytes 
cells of the cartilage 


collagen fiber 
flexible fibrous proteins that give connective tissue tensile strength 


connective tissue proper 
connective tissue containing a viscous matrix, fibers, and cells. 


dense connective tissue 
connective tissue proper that contains many fibers that provide both 
elasticity and protection 


elastic cartilage 
type of cartilage, with elastin as the major protein, characterized by 
rigid support as well as elasticity 


elastic fiber 
fibrous protein within connective tissue that contains a high percentage 
of the protein elastin that allows the fibers to stretch and return to 
original size 


fibroblast 
most abundant cell type in connective tissue, secretes protein fibers 
and matrix into the extracellular space 


fibrocartilage 
tough form of cartilage, made of thick bundles of collagen fibers 
embedded in chondroitin sulfate ground substance 


fibrocyte 
less active form of fibroblast 


fluid connective tissue 
specialized cells that circulate in a watery fluid containing salts, 
nutrients, and dissolved proteins 


ground substance 
fluid or semi-fluid portion of the matrix 


hyaline cartilage 
most common type of cartilage, smooth and made of short collagen 
fibers embedded in a chondroitin sulfate ground substance 


lacunae 
(singular = lacuna) small spaces in bone or cartilage tissue that cells 
occupy 


loose connective tissue 
(also, areolar tissue) type of connective tissue proper that shows little 
specialization with cells dispersed in the matrix 


matrix 
extracellular material which is produced by the cells embedded in it, 
containing ground substance and fibers 


mesenchymal cell 
adult stem cell from which most connective tissue cells are derived 


mesenchyme 
embryonic tissue from which connective tissue cells derive 


mucous connective tissue 
specialized loose connective tissue present in the umbilical cord 


parenchyma 
functional cells of a gland or organ, in contrast with the supportive or 
connective tissue of a gland or organ 


reticular fiber 
fine fibrous protein, made of collagen subunits, which cross-link to 
form supporting “nets” within connective tissue 


reticular tissue 
type of loose connective tissue that provides a supportive framework 
to soft organs, such as lymphatic tissue, spleen, and the liver 


supportive connective tissue 
type of connective tissue that provides strength to the body and 
protects soft tissue 


Muscle Tissue and Motion 
By the end of this section, you will be able to: 


e Identify the three types of muscle tissue 
¢ Compare and contrast the functions of each muscle tissue type 
e Explain how muscle tissue can enable motion 


Muscle tissue is characterized by properties that allow movement. Muscle 
cells are excitable; they respond to a stimulus. They are contractile, 
meaning they can shorten and generate a pulling force. When attached 
between two movable objects, in other words, bones, contractions of the 
muscles cause the bones to move. Some muscle movement is voluntary, 


which means it is under conscious control. For example, a person decides to 


open a book and read a chapter on anatomy. Other movements are 
involuntary, meaning they are not under conscious control, such as the 
contraction of your pupil in bright light. Muscle tissue is classified into 
three types according to structure and function: skeletal, cardiac, and 


smooth ({link]). 


Comparison of Structure and Properties of Muscle Tissue Types 


Tissue 


Skeletal 


Histology 


Long 
cylindrical 
fiber, 
Striated, 
many 
peripherally 
located 
nuclei 


Function 


Voluntary 
movement, produces 
heat, protects organs 


Location 


Attached to 
bones and 
around 
entrance 
points to 
body (e.g., 
mouth, 
anus) 


Comparison of Structure and Properties of Muscle Tissue Types 


Tissue Histology Function Location 
Short, 
branched, 
: rl ntr m 
aie st ated, Contracts to pump ean 
single blood 
central 
nucleus 
Involuntar 
Short, vey 
: movement, moves 
ae food, involuntar 
shaped, no ‘ y Walls of 
: control of : 
evident Pea major 
Smooth ihe respiration, moves 
Striation, organs and 
single secretions, regulates passageways 
F flow of blood in 
SSA eae arteries b 
each fiber y 


contraction 


Skeletal muscle is attached to bones and its contraction makes possible 
locomotion, facial expressions, posture, and other voluntary movements of 
the body. Forty percent of your body mass is made up of skeletal muscle. 
Skeletal muscles generate heat as a byproduct of their contraction and thus 
participate in thermal homeostasis. Shivering is an involuntary contraction 
of skeletal muscles in response to perceived lower than normal body 
temperature. The muscle cell, or myocyte, develops from myoblasts 
derived from the mesoderm. Myocytes and their numbers remain relatively 
constant throughout life. Skeletal muscle tissue is arranged in bundles 
surrounded by connective tissue. Under the light microscope, muscle cells 
appear striated with many nuclei squeezed along the membranes. The 
striation is due to the regular alternation of the contractile proteins actin 
and myosin, along with the structural proteins that couple the contractile 
proteins to connective tissues. The cells are multinucleated as a result of the 
fusion of the many myoblasts that fuse to form each long muscle fiber. 


Cardiac muscle forms the contractile walls of the heart. The cells of 
cardiac muscle, known as cardiomyocytes, also appear striated under the 
microscope. Unlike skeletal muscle fibers, cardiomyocytes are single cells 
typically with a single centrally located nucleus. A principal characteristic 
of cardiomyocytes is that they contract on their own intrinsic rhythms 
without any external stimulation. Cardiomyocyte attach to one another with 
specialized cell junctions called intercalated discs. Intercalated discs have 
both anchoring junctions and gap junctions. Attached cells form long, 
branching cardiac muscle fibers that are, essentially, a mechanical and 
electrochemical syncytium allowing the cells to synchronize their actions. 
The cardiac muscle pumps blood through the body and is under involuntary 
control. The attachment junctions hold adjacent cells together across the 
dynamic pressures changes of the cardiac cycle. 


Smooth muscle tissue contraction is responsible for involuntary 
movements in the internal organs. It forms the contractile component of the 
digestive, urinary, and reproductive systems as well as the airways and 
arteries. Each cell is spindle shaped with a single nucleus and no visible 
striations ((link]). 

Muscle Tissue 


(a) Skeletal muscle cells have 
prominent striation and 
nuclei on their periphery. (b) 
Smooth muscle cells have a 
single nucleus and no visible 
striations. (c) Cardiac muscle 
cells appear striated and have 
a single nucleus. From top, 
LM x 1600, LM x 1600, LM 
x 1600. (Micrographs 
provided by the Regents of 
University of Michigan 
Medical School © 2012) 


Note: 


| acl 
ele, "a 


—: 
mess Openstax COLLEGE 
—— . 

. 


Watch this video to learn more about muscle tissue. In looking through a 
microscope how could you distinguish skeletal muscle tissue from smooth 
muscle? 


Chapter Review 


The three types of muscle cells are skeletal, cardiac, and smooth. Their 
morphologies match their specific functions in the body. Skeletal muscle is 
voluntary and responds to conscious stimuli. The cells are striated and 
multinucleated appearing as long, unbranched cylinders. Cardiac muscle is 
involuntary and found only in the heart. Each cell is striated with a single 
nucleus and they attach to one another to form long fibers. Cells are 
attached to one another at intercalated disks. The cells are interconnected 
physically and electrochemically to act as a syncytium. Cardiac muscle 
cells contract autonomously and involuntarily. Smooth muscle is 
involuntary. Each cell is a spindle-shaped fiber and contains a single 
nucleus. No striations are evident because the actin and myosin filaments 
do not align in the cytoplasm. 


Interactive Link Questions 


Exercise: 
Problem: 
Watch this video to learn more about muscle tissue. In looking through 


a microscope how could you distinguish skeletal muscle tissue from 
smooth muscle? 


Solution: 


Skeletal muscle cells are striated. 


Review Questions 
Exercise: 
Problem: 


Striations, cylindrical cells, and multiple nuclei are observed in 


a. Skeletal muscle only 
b. cardiac muscle only 
c. smooth muscle only 
d. skeletal and cardiac muscles 


Solution: 


A 


Exercise: 


Problem:The cells of muscles, myocytes, develop from 


a. myoblasts 
b. endoderm 
c. fibrocytes 
d. chondrocytes 


Solution: 


A 


Exercise: 


Problem: 


Skeletal muscle is composed of very hard working cells. Which 
organelles do you expect to find in abundance in skeletal muscle cell? 


a. nuclei 

b. striations 

c. golgi bodies 
d. mitochondria 


Solution: 


D 


Critical Thinking Questions 


Exercise: 
Problem: 
You are watching cells in a dish spontaneously contract. They are all 
contracting at different rates; some fast, some slow. After a while, 


several cells link up and they begin contracting in synchrony. Discuss 
what is going on and what type of cells you are looking at. 


Solution: 
The cells in the dish are cardiomyocytes, cardiac muscle cells. They 
have an intrinsic ability to contract. When they link up, they form 


intercalating discs that allow the cells to communicate with each other 
and begin contracting in synchrony. 


Exercise: 


Problem: Why does skeletal muscle look striated? 


Solution: 


Under the light microscope, cells appear striated due to the 
arrangement of the contractile proteins actin and myosin. 


Glossary 


cardiac muscle 
heart muscle, under involuntary control, composed of striated cells that 
attach to form fibers, each cell contains a single nucleus, contracts 
autonomously 


myocyte 
muscle cells 


skeletal muscle 
usually attached to bone, under voluntary control, each cell is a fiber 
that is multinucleated and striated 


smooth muscle 
under involuntary control, moves internal organs, cells contain a single 
nucleus, are spindle-shaped, and do not appear striated; each cell is a 
fiber 


striation 
alignment of parallel actin and myosin filaments which form a banded 
pattern 


Nervous Tissue Mediates Perception and Response 
By the end of this section, you will be able to: 


e Identify the classes of cells that make up nervous tissue 
e Discuss how nervous tissue mediates perception and response 


Nervous tissue is characterized as being excitable and capable of sending 
and receiving electrochemical signals that provide the body with 
information. Two main classes of cells make up nervous tissue: the neuron 
and neuroglia ([link]). Neurons propagate information via electrochemical 
impulses, called action potentials, which are biochemically linked to the 
release of chemical signals. Neuroglia play an essential role in supporting 
neurons and modulating their information propagation. 

The Neuron 


Contact with 
other cells 


a Nucleus 
Microfibrils and 


microtubules 


Axon 


Dendrites 


The cell body of a neuron, also called the soma, 
contains the nucleus and mitochondria. The dendrites 
transfer the nerve impulse to the soma. The axon 
carries the action potential away to another excitable 
cell. LM x 1600. (Micrograph provided by the 
Regents of University of Michigan Medical School © 
2012) 


Note: 


= 
mss Openstax COLLEGE 


Follow this link to learn more about nervous tissue. What are the main 
parts of a nerve cell? 


Neurons display distinctive morphology, well suited to their role as 
conducting cells, with three main parts. The cell body includes most of the 
cytoplasm, the organelles, and the nucleus. Dendrites branch off the cell 
body and appear as thin extensions. A long “tail,” the axon, extends from 
the neuron body and can be wrapped in an insulating layer known as 
myelin, which is formed by accessory cells. The synapse is the gap between 
nerve cells, or between a nerve cell and its target, for example, a muscle or 
a gland, across which the impulse is transmitted by chemical compounds 
known as neurotransmitters. Neurons categorized as multipolar neurons 
have several dendrites and a single prominent axon. Bipolar neurons 
possess a single dendrite and axon with the cell body, while unipolar 
neurons have only a single process extending out from the cell body, which 
divides into a functional dendrite and into a functional axon. When a neuron 
is sufficiently stimulated, it generates an action potential that propagates 
down the axon towards the synapse. If enough neurotransmitters are 
released at the synapse to stimulate the next neuron or target, a response is 
generated. 


The second class of neural cells comprises the neuroglia or glial cells, 
which have been characterized as having a simple support role. The word 
“glia” comes from the Greek word for glue. Recent research is shedding 
light on the more complex role of neuroglia in the function of the brain and 
nervous system. Astrocyte cells, named for their distinctive star shape, are 
abundant in the central nervous system. The astrocytes have many 


functions, including regulation of ion concentration in the intercellular 
space, uptake and/or breakdown of some neurotransmitters, and formation 
of the blood-brain barrier, the membrane that separates the circulatory 
system from the brain. Microglia protect the nervous system against 
infection but are not nervous tissue because they are related to 
macrophages. Oligodendrocyte cells produce myelin in the central nervous 
system (brain and spinal cord) while the Schwann cell produces myelin in 
the peripheral nervous system ([link]). 

Nervous Tissue 


Neurons 


Microglial cell Sa Astrocytes y 


' “s 
aK. OZ 


Oligodendrocyte 


Nervous tissue is made up of neurons and neuroglia. The cells 
of nervous tissue are specialized to transmit and receive 
impulses. LM x 872. (Micrograph provided by the Regents of 
University of Michigan Medical School © 2012) 


Chapter Review 


The most prominent cell of the nervous tissue, the neuron, is characterized 
mainly by its ability to receive stimuli and respond by generating an 
electrical signal, known as an action potential, which can travel rapidly over 
great distances in the body. A typical neuron displays a distinctive 
morphology: a large cell body branches out into short extensions called 
dendrites, which receive chemical signals from other neurons, and a long 
tail called an axon, which relays signals away from the cell to other 
neurons, muscles, or glands. Many axons are wrapped by a myelin sheath, a 


lipid derivative that acts as an insulator and speeds up the transmission of 
the action potential. Other cells in the nervous tissue, the neuroglia, include 
the astrocytes, microglia, oligodendrocytes, and Schwann cells. 


Interactive Link Questions 


Exercise: 


Problem: 


Follow this link to learn more about nervous tissue. What are the main 
parts of a nerve cell? 


Solution: 


Dendrites, cell body, and the axon. 


Review Questions 


Exercise: 


Problem: 


The cells responsible for the transmission of the nerve impulse are 


a. Neurons 
b. oligodendrocytes 
c. astrocytes 
d. microglia 


Solution: 


A 


Exercise: 


Problem: 


The nerve impulse travels down a(n) , away from the cell 
body. 


a. dendrite 

b. axon 

c. microglia 

d. collagen fiber 


Solution: 


B 
Exercise: 


Problem: 


Which of the following central nervous system cells regulate ions, 
regulate the uptake and/or breakdown of some neurotransmitters, and 
contribute to the formation of the blood-brain barrier? 


a. microglia 
b. neuroglia 
c. oligodendrocytes 
d. astrocytes 


Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: 


Which morphological adaptations of neurons make them suitable for 
the transmission of nerve impulse? 


Solution: 
Neurons are well suited for the transmission of nerve impulses because 
short extensions, dendrites, receive impulses from other neurons, while 


a long tail extension, an axon, carries electrical impulses away from 
the cell to other neurons. 


Exercise: 
Problem: What are the functions of astrocytes? 


Solution: 

Astrocytes regulate ions and uptake and/or breakdown of some 
neurotransmitters and contribute to the formation of the blood-brain- 
barrier. 


References 


Stern, P. Focus issue: getting excited about glia. Science [Internet]. 2010 
[cited 2012 Dec 4]; 3(147):330-773. Available from: 


http://stke.sciencemag,org/cgi/content/abstract/sigtrans;3/147/eg11 


Ming GL, Song H. Adult neurogenesis in the mammalian central nervous 
system. Annu. Rev. Neurosci. 2005 [cited 2012 Dec 4]; 28:223—250. 


Glossary 


astrocyte 


star-shaped cell in the central nervous system that regulates ions and 
uptake and/or breakdown of some neurotransmitters and contributes to 
the formation of the blood-brain barrier 


myelin 
layer of lipid inside some neuroglial cells that wraps around the axons 
of some neurons 


neuroglia 
supportive neural cells 


neuron 
excitable neural cell that transfer nerve impulses 


oligodendrocyte 
neuroglial cell that produces myelin in the brain 


Schwann cell 
neuroglial cell that produces myelin in the peripheral nervous system 


Tissue Injury and Aging 
By the end of this section, you will be able to: 


e Identify the cardinal signs of inflammation 

List the body’s response to tissue injury 

e Explain the process of tissue repair 

Discuss the progressive impact of aging on tissue 
e Describe cancerous mutations’ effect on tissue 


Tissues of all types are vulnerable to injury and, inevitably, aging. In the 
former case, understanding how tissues respond to damage can guide 
strategies to aid repair. In the latter case, understanding the impact of aging 
can help in the search for ways to diminish its effects. 


Tissue Injury and Repair 


Inflammation is the standard, initial response of the body to injury. 
Whether biological, chemical, physical, or radiation burns, all injuries lead 
to the same sequence of physiological events. Inflammation limits the 
extent of injury, partially or fully eliminates the cause of injury, and initiates 
repair and regeneration of damaged tissue. Necrosis, or accidental cell 
death, causes inflammation. Apoptosis is programmed cell death, a normal 
step-by-step process that destroys cells no longer needed by the body. By 
mechanisms still under investigation, apoptosis does not initiate the 
inflammatory response. Acute inflammation resolves over time by the 
healing of tissue. If inflammation persists, it becomes chronic and leads to 
diseased conditions. Arthritis and tuberculosis are examples of chronic 
inflammation. The suffix “-itis” denotes inflammation of a specific organ or 
type, for example, peritonitis is the inflammation of the peritoneum, and 
meningitis refers to the inflammation of the meninges, the tough 
membranes that surround the central nervous system 


The four cardinal signs of inflammation—redness, swelling, pain, and local 
heat—were first recorded in antiquity. Cornelius Celsus is credited with 
documenting these signs during the days of the Roman Empire, as early as 
the first century AD. A fifth sign, loss of function, may also accompany 
inflammation. 


Upon tissue injury, damaged cells release inflammatory chemical signals 
that evoke local vasodilation, the widening of the blood vessels. Increased 
blood flow results in apparent redness and heat. In response to injury, mast 
cells present in tissue degranulate, releasing the potent vasodilator 
histamine. Increased blood flow and inflammatory mediators recruit white 
blood cells to the site of inflammation. The endothelium lining the local 
blood vessel becomes “leaky” under the influence of histamine and other 
inflammatory mediators allowing neutrophils, macrophages, and fluid to 
move from the blood into the interstitial tissue spaces. The excess liquid in 
tissue causes swelling, more properly called edema. The swollen tissues 
squeezing pain receptors cause the sensation of pain. Prostaglandins 
released from injured cells also activate pain neurons. Non-steroidal anti- 
inflammatory drugs (NSAIDs) reduce pain because they inhibit the 
synthesis of prostaglandins. High levels of NSAIDs reduce inflammation. 
Antihistamines decrease allergies by blocking histamine receptors and as a 
result the histamine response. 


After containment of an injury, the tissue repair phase starts with removal of 
toxins and waste products. Clotting (coagulation) reduces blood loss from 
damaged blood vessels and forms a network of fibrin proteins that trap 
blood cells and bind the edges of the wound together. A scab forms when 
the clot dries, reducing the risk of infection. Sometimes a mixture of dead 
leukocytes and fluid called pus accumulates in the wound. As healing 
progresses, fibroblasts from the surrounding connective tissues replace the 
collagen and extracellular material lost by the injury. Angiogenesis, the 
growth of new blood vessels, results in vascularization of the new tissue 
known as granulation tissue. The clot retracts pulling the edges of the 
wound together, and it slowly dissolves as the tissue is repaired. When a 
large amount of granulation tissue forms and capillaries disappear, a pale 
scar is often visible in the healed area. A primary union describes the 
healing of a wound where the edges are close together. When there is a 
gaping wound, it takes longer to refill the area with cells and collagen. The 
process called secondary union occurs as the edges of the wound are 
pulled together by what is called wound contraction. When a wound is 
more than one quarter of an inch deep, sutures (stitches) are recommended 
to promote a primary union and avoid the formation of a disfiguring scar. 


Regeneration is the addition of new cells of the same type as the ones that 
were injured ((link]). 
Tissue Healing 


Clotting occurs, caused by clotting proteins Epithelial cells multiply and fill Restored epthelium thickens; the 
and plasma proteins, and a scab is formed in over the granulation tissue area matures and contracts 


Inflammatory chemicals White blood cells seep Granulation tissue restores Underlying area 
are released from injury _ into the injured area the vascular supply of scar tissue 


During wound repair, collagen fibers are laid down 
randomly by fibroblasts that move into repair the area. 


Note: 


Oa ec 


openstax COLLEGE” 
. 75 

. 
O) 
a r 


Watch this video to see a hand heal. Over what period of time do you think 
these images were taken? 


Tissue and Aging 


According to poet Ralph Waldo Emerson, “The surest poison is time.” In 
fact, biology confirms that many functions of the body decline with age. All 
the cells, tissues, and organs are affected by senescence, with noticeable 
variability between individuals owing to different genetic makeup and 
lifestyles. The outward signs of aging are easily recognizable. The skin and 
other tissues become thinner and drier, reducing their elasticity, contributing 
to wrinkles and high blood pressure. Hair turns gray because follicles 
produce less melanin, the brown pigment of hair and the iris of the eye. The 
face looks flabby because elastic and collagen fibers decrease in connective 
tissue and muscle tone is lost. Glasses and hearing aids may become parts 
of life as the senses slowly deteriorate, all due to reduced elasticity. Overall 
height decreases as the bones lose calcium and other minerals. With age, 
fluid decreases in the fibrous cartilage disks intercalated between the 
vertebrae in the spine. Joints lose cartilage and stiffen. Many tissues, 
including those in muscles, lose mass through a process called atrophy. 
Lumps and rigidity become more widespread. As a consequence, the 
passageways, blood vessels, and airways become more rigid. The brain and 
spinal cord lose mass. Nerves do not transmit impulses with the same speed 
and frequency as in the past. Some loss of thought clarity and memory can 
accompany aging. More severe problems are not necessarily associated 
with the aging process and may be symptoms of underlying illness. 


As exterior signs of aging increase, so do the interior signs, which are not as 
noticeable. The incidence of heart diseases, respiratory syndromes, and type 
2 diabetes increases with age, though these are not necessarily age- 
dependent effects. Wound healing is slower in the elderly, accompanied by 
a higher frequency of infection as the capacity of the immune system to 
fend off pathogen declines. 


Aging is also apparent at the cellular level because all cells experience 
changes with aging. Telomeres, regions of the chromosomes necessary for 
cell division, shorten each time cells divide. As they do, cells are less able 
to divide and regenerate. Because of alterations in cell membranes, 
transport of oxygen and nutrients into the cell and removal of carbon 
dioxide and waste products from the cell are not as efficient in the elderly. 
Cells may begin to function abnormally, which may lead to diseases 
associated with aging, including arthritis, memory issues, and some cancers. 


The progressive impact of aging on the body varies considerably among 
individuals, but Studies indicate, however, that exercise and healthy 
lifestyle choices can slow down the deterioration of the body that comes 
with old age. 


Note: 

Homeostatic Imbalances 

Tissues and Cancer 

Cancer is a generic term for many diseases in which cells escape regulatory 
signals. Uncontrolled growth, invasion into adjacent tissues, and 
colonization of other organs, if not treated early enough, are its hallmarks. 
Health suffers when tumors “rob” blood supply from the “normal” organs. 
A mutation is defined as a permanent change in the DNA of a cell. 
Epigenetic modifications, changes that do not affect the code of the DNA 
but alter how the DNA is decoded, are also known to generate abnormal 
cells. Alterations in the genetic material may be caused by environmental 
agents, infectious agents, or errors in the replication of DNA that 
accumulate with age. Many mutations do not cause any noticeable change 
in the functions of a cell. However, if the modification affects key proteins 
that have an impact on the cell’s ability to proliferate in an orderly fashion, 
the cell starts to divide abnormally. As changes in cells accumulate, they 
lose their ability to form regular tissues. A tumor, a mass of cells 
displaying abnormal architecture, forms in the tissue. Many tumors are 
benign, meaning they do not metastasize nor cause disease. A tumor 
becomes malignant, or cancerous, when it breaches the confines of its 
tissue, promotes angiogenesis, attracts the growth of capillaries, and 
metastasizes to other organs ((link]). The specific names of cancers reflect 
the tissue of origin. Cancers derived from epithelial cells are referred to as 
carcinomas. Cancer in myeloid tissue or blood cells form myelomas. 
Leukemias are cancers of white blood cells, whereas sarcomas derive from 
connective tissue. Cells in tumors differ both in structure and function. 
Some cells, called cancer stem cells, appear to be a subtype of cell 
responsible for uncontrolled growth. Recent research shows that contrary 
to what was previously assumed, tumors are not disorganized masses of 
cells, but have their own structures. 


Development of Cancer 


Cell division 
takes place 
to replace 
lost tissue 
Cell division 
accelerates 


.—\— 


WSS 
SS 
SW: 
nN 


Carcinoma 
breaks into 
underlying 
tissue 
Underlying 
tissue 


Note the change in cell size, 
nucleus size, and organization 


in the tissue. 


Note: 


Coker aa) 


ae 


— 
wm, OPENSTAX COLLEGE 
— 

. 


- 


tO) hs 


Watch this video to learn more about tumors. What is a tumor? 


Cancer treatments vary depending on the disease’s type and stage. 
Traditional approaches, including surgery, radiation, chemotherapy, and 
hormonal therapy, aim to remove or kill rapidly dividing cancer cells, but 
these strategies have their limitations. Depending on a tumor’s location, for 
example, cancer surgeons may be unable to remove it. Radiation and 
chemotherapy are difficult, and it is often impossible to target only the 
cancer cells. The treatments inevitably destroy healthy tissue as well. To 
address this, researchers are working on pharmaceuticals that can target 
specific proteins implicated in cancer-associated molecular pathways. 


Chapter Review 


Inflammation is the classic response of the body to injury and follows a 
common sequence of events. The area is red, feels warm to the touch, 
swells, and is painful. Injured cells, mast cells, and resident macrophages 
release chemical signals that cause vasodilation and fluid leakage in the 
surrounding tissue. The repair phase includes blood clotting, followed by 
regeneration of tissue as fibroblasts deposit collagen. Some tissues 
regenerate more readily than others. Epithelial and connective tissues 
replace damaged or dead cells from a supply of adult stem cells. Muscle 
and nervous tissues undergo either slow regeneration or do not repair at all. 


Age affects all the tissues and organs of the body. Damaged cells do not 
regenerate as rapidly as in younger people. Perception of sensation and 
effectiveness of response are lost in the nervous system. Muscles atrophy, 


and bones lose mass and become brittle. Collagen decreases in some 
connective tissue, and joints stiffen. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to see a hand heal. Over what period of time do you 
think these images were taken? 


Solution: 


Approximately one month. 
Exercise: 


Problem: 
Watch this video to learn more about tumors. What is a tumor? 
Solution: 


A mass of cancer cells that continue to grow and divide. 


Review Questions 


Exercise: 


Problem: 


Which of the following processes is not a cardinal sign of 
inflammation? 


a. redness 
b. heat 

c. fever 

d. swelling 


Solution: 


C 
Exercise: 


Problem: 


When a mast cell reacts to an irritation, which of the following 
chemicals does it release? 


a. collagen 

b. histamine 

c. hyaluronic acid 
d. meylin 


Solution: 


B 


Exercise: 


Problem: Atrophy refers to 


a. loss of elasticity 

b. loss of mass 

c. loss of rigidity 

d. loss of permeability 


Solution: 


B 


Exercise: 


Problem: 


Individuals can slow the rate of aging by modifying all of these 
lifestyle aspects except for 


a. diet 

b. exercise 

c. genetic factors 
d. stress 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: 


Why is it important to watch for increased redness, swelling and pain 
after a cut or abrasion has been cleaned and bandaged? 


Solution: 


These symptoms would indicate that infection is present. 


Exercise: 


Problem: 


Aspirin is a non-steroidal anti-inflammatory drug (NSAID) that 
inhibits the formation of blood clots and is taken regularly by 
individuals with a heart condition. Steroids such as cortisol are used to 
control some autoimmune diseases and severe arthritis by down- 
regulating the inflammatory response. After reading the role of 
inflammation in the body’s response to infection, can you predict an 
undesirable consequence of taking anti-inflammatory drugs on a 
regular basis? 


Solution: 


Since NSAIDs or other anti-inflammatory drugs inhibit the formation 
of blood clots, regular and prolonged use of these drugs may promote 
internal bleeding, such as bleeding in the stomach. Excessive levels of 
cortisol would suppress inflammation, which could slow the wound 
healing process. 


Exercise: 
Problem: 
As an individual ages, a constellation of symptoms begins the decline 
to the point where an individual’s functioning is compromised. 


Identify and discuss two factors that have a role in factors leading to 
the compromised situation. 


Solution: 
The genetic makeup and the lifestyle of each individual are factors 


which determine the degree of decline in cells, tissues, and organs as 
an individual ages. 


Exercise: 


Problem: Discuss changes that occur in cells as a person ages. 


Solution: 


All cells experience changes with aging. They become larger, and 
many cannot divide and regenerate. Because of alterations in cell 
membranes, transport of oxygen and nutrients into the cell and 
removal of carbon dioxide and waste products are not as efficient in 
the elderly. Cells lose their ability to function, or they begin to function 
abnormally, leading to disease and cancer. 


References 


Emerson, RW. Old age. Atlantic. 1862 [cited 2012 Dec 4]; 9(51):134—140. 


Glossary 


apoptosis 
programmed cell death 


atrophy 
loss of mass and function 


clotting 
also called coagulation; complex process by which blood components 
form a plug to stop bleeding 


histamine 
chemical compound released by mast cells in response to injury that 
causes vasodilation and endothelium permeability 


inflammation 
response of tissue to injury 


necrosis 
accidental death of cells and tissues 


primary union 
condition of a wound where the wound edges are close enough to be 
brought together and fastened if necessary, allowing quicker and more 
thorough healing 


secondary union 
wound healing facilitated by wound contraction 


vasodilation 
widening of blood vessels 


wound contraction 
process whereby the borders of a wound are physically drawn together 


Layers of the Skin 
By the end of this section, you will be able to: 


e Identify the components of the integumentary system 
Describe the layers of the skin and the functions of each layer 
e Identify and describe the hypodermis and deep fascia 
Describe the role of keratinocytes and their life cycle 

e Describe the role of melanocytes in skin pigmentation 


Although you may not typically think of the skin as an organ, it is in fact 
made of tissues that work together as a single structure to perform unique 
and critical functions. The skin and its accessory structures make up the 
integumentary system, which provides the body with overall protection. 
The skin is made of multiple layers of cells and tissues, which are held to 
underlying structures by connective tissue ({link]). The deeper layer of skin 
is well vascularized (has numerous blood vessels). It also has numerous 
sensory, and autonomic and sympathetic nerve fibers ensuring 
communication to and from the brain. 

Layers of Skin 


Hair shaft 
Pore of sweat gland duct 


Epidermis 


| 


Arrector pili 
muscle 


Hair follicle 


Sebaceous (oil) 
gland 


Hypodermis 
Hair root 


Hair follicle 


receptor Eccrine sweat gland 
Pacinian corpuscle 


Cutaneous vascular 
plexus 


Adipose tissue 


Sensory nerve fiber 


The skin is composed of two main layers: the epidermis, 
made of closely packed epithelial cells, and the dermis, 
made of dense, irregular connective tissue that houses 
blood vessels, hair follicles, sweat glands, and other 
structures. Beneath the dermis lies the hypodermis, 
which is composed mainly of loose connective and fatty 
tissues. 


ee 


mess’ OPENStAX COLLEGE” 
i 
io ae T 


The skin consists of two main layers and a closely associated layer. View 
this animation to learn more about layers of the skin. What are the basic 
functions of each of these layers? 


The Epidermis 


The epidermis is composed of keratinized, stratified squamous epithelium. 
It is made of four or five layers of epithelial cells, depending on its location 
in the body. It does not have any blood vessels within it (i.e., it is 
avascular). Skin that has four layers of cells is referred to as “thin skin.” 
From deep to superficial, these layers are the stratum basale, stratum 
spinosum, stratum granulosum, and stratum corneum. Most of the skin can 
be classified as thin skin. “Thick skin” is found only on the palms of the 
hands and the soles of the feet. It has a fifth layer, called the stratum 


lucidum, located between the stratum corneum and the stratum granulosum 
(Llink]). 
Thin Skin versus Thick Skin 


These slides show cross- 
sections of the epidermis and 
dermis of (a) thin and (b) 
thick skin. Note the 
significant difference in the 
thickness of the epithelial 
layer of the thick skin. From 
top, LM x 40, LM ~x 40. 
(Micrographs provided by the 
Regents of University of 
Michigan Medical School © 
2012) 


The cells in all of the layers except the stratum basale are called 
keratinocytes. A keratinocyte is a cell that manufactures and stores the 
protein keratin. Keratin is an intracellular fibrous protein that gives hair, 
nails, and skin their hardness and water-resistant properties. The 
keratinocytes in the stratum corneum are dead and regularly slough away, 
being replaced by cells from the deeper layers ({link]). 

Epidermis 


The epidermis is epithelium 
composed of multiple layers 
of cells. The basal layer 
consists of cuboidal cells, 
whereas the outer layers are 
squamous, keratinized cells, 
so the whole epithelium is 
often described as being 
keratinized stratified 
squamous epithelium. LM x 
40. (Micrograph provided by 
the Regents of University of 
Michigan Medical School © 
2012) 


Note: 


. . 
openstax COLLEGE 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. If you zoom on the cells at the outermost layer of this 
section of skin, what do you notice about the cells? 


Stratum Basale 


The stratum basale (also called the stratum germinativum) is the deepest 
epidermal layer and attaches the epidermis to the basal lamina, below which 
lie the layers of the dermis. The cells in the stratum basale bond to the 
dermis via intertwining collagen fibers, referred to as the basement 
membrane. A finger-like projection, or fold, known as the dermal papilla 
(plural = dermal papillae) is found in the superficial portion of the dermis. 
Dermal papillae increase the strength of the connection between the 
epidermis and dermis; the greater the folding, the stronger the connections 
made ([link]). 

Layers of the Epidermis 


Dead cells filled 
with keratin 


5 = 
Stratum corneum = 
eS =o SSSqNqBPHOO 


Stratum lucidum ——_{_ 


Stratum granulosum 


: ——_,_ 
Stratum spinosum = 


Stratum basale — j 


Melanocyte 


Lamellar granules 


Keratinocyte 


Merkel cell 


: Sensory neuron 
Dermis 


The epidermis of thick skin has five layers: stratum 
basale, stratum spinosum, stratum granulosum, stratum 
lucidum, and stratum corneum. 


The stratum basale is a single layer of cells primarily made of basal cells. A 
basal cell is a cuboidal-shaped stem cell that is a precursor of the 
keratinocytes of the epidermis. All of the keratinocytes are produced from 
this single layer of cells, which are constantly going through mitosis to 
produce new cells. As new cells are formed, the existing cells are pushed 
superficially away from the stratum basale. Two other cell types are found 
dispersed among the basal cells in the stratum basale. The first is a Merkel 
cell, which functions as a receptor and is responsible for stimulating 
sensory nerves that the brain perceives as touch. These cells are especially 
abundant on the surfaces of the hands and feet. The second is a melanocyte, 
a cell that produces the pigment melanin. Melanin gives hair and skin its 
color, and also helps protect the living cells of the epidermis from 
ultraviolet (UV) radiation damage. 


In a growing fetus, fingerprints form where the cells of the stratum basale 
meet the papillae of the underlying dermal layer (papillary layer), resulting 
in the formation of the ridges on your fingers that you recognize as 
fingerprints. Fingerprints are unique to each individual and are used for 
forensic analyses because the patterns do not change with the growth and 
aging processes. 


Stratum Spinosum 


As the name suggests, the stratum spinosum is spiny in appearance due to 
the protruding cell processes that join the cells via a structure called a 
desmosome. The desmosomes interlock with each other and strengthen the 
bond between the cells. It is interesting to note that the “spiny” nature of 
this layer is an artifact of the staining process. Unstained epidermis samples 
do not exhibit this characteristic appearance. The stratum spinosum is 
composed of eight to 10 layers of keratinocytes, formed as a result of cell 
division in the stratum basale ([link]). Interspersed among the keratinocytes 
of this layer is a type of dendritic cell called the Langerhans cell, which 
functions as a macrophage by engulfing bacteria, foreign particles, and 
damaged cells that occur in this layer. 

Cells of the Epidermis 


065 - Epidermis_001.svs 


WebScope 1 
16013 x 16013 size 733.61MB mag 20X Return to image directory 
(© fale "SSB Oo Gm O OVE | 
Epiaermis 
Stratum ,, 
corneum i 
Stratum ~ 
granulosum 
Remnants of 
cross-sectioned 
Stratum shed hair and 
spinosum its follicle 
Stratum basal 
or germinativum 
Capillai 
Dermi poy 
EJM Mag. 2,700 X 


The cells in the different layers of the epidermis originate 
from basal cells located in the stratum basale, yet the 
cells of each layer are distinctively different. EM x 2700. 
(Micrograph provided by the Regents of University of 
Michigan Medical School © 2012) 


Note: 


poe 


openstax COLLEGE” 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. If you zoom on the cells at the outermost layer of this 
section of skin, what do you notice about the cells? 


The keratinocytes in the stratum spinosum begin the synthesis of keratin 
and release a water-repelling glycolipid that helps prevent water loss from 
the body, making the skin relatively waterproof. As new keratinocytes are 
produced atop the stratum basale, the keratinocytes of the stratum spinosum 
are pushed into the stratum granulosum. 


Stratum Granulosum 


The stratum granulosum has a grainy appearance due to further changes 
to the keratinocytes as they are pushed from the stratum spinosum. The 
cells (three to five layers deep) become flatter, their cell membranes 
thicken, and they generate large amounts of the proteins keratin, which is 
fibrous, and keratohyalin, which accumulates as lamellar granules within 
the cells (see [link]). These two proteins make up the bulk of the 
keratinocyte mass in the stratum granulosum and give the layer its grainy 
appearance. The nuclei and other cell organelles disintegrate as the cells 
die, leaving behind the keratin, keratohyalin, and cell membranes that will 
form the stratum lucidum, the stratum comeum, and the accessory 
structures of hair and nails. 


Stratum Lucidum 


The stratum lucidum is a smooth, seemingly translucent layer of the 
epidermis located just above the stratum granulosum and below the stratum 
corneum. This thin layer of cells is found only in the thick skin of the 
palms, soles, and digits. The keratinocytes that compose the stratum 
lucidum are dead and flattened (see [link]). These cells are densely packed 
with eleiden, a clear protein rich in lipids, derived from keratohyalin, which 


gives these cells their transparent (i.e., lucid) appearance and provides a 
barrier to water. 


Stratum Corneum 


The stratum corneum is the most superficial layer of the epidermis and is 
the layer exposed to the outside environment (see [link]). The increased 
keratinization (also called cornification) of the cells in this layer gives it its 
name. There are usually 15 to 30 layers of cells in the stratum corneum. 
This dry, dead layer helps prevent the penetration of microbes and the 
dehydration of underlying tissues, and provides a mechanical protection 
against abrasion for the more delicate, underlying layers. Cells in this layer 
are shed periodically and are replaced by cells pushed up from the stratum 
granulosum (or stratum lucidum in the case of the palms and soles of feet). 
The entire layer is replaced during a period of about 4 weeks. Cosmetic 
procedures, such as microdermabrasion, help remove some of the dry, upper 
layer and aim to keep the skin looking “fresh” and healthy. 


Dermis 


The dermis might be considered the “core” of the integumentary system 
(derma- = “skin”), as distinct from the epidermis (epi- = “upon” or “over” 
and hypodermis (hypo- = “below”). It contains blood and lymph vessels, 
nerves, and other structures, such as hair follicles and sweat glands. The 
dermis is made of two layers of connective tissue that compose an 
interconnected mesh of elastin and collagenous fibers, produced by 
fibroblasts ({link]). 


Layers of the Dermis 


This stained slide shows the two 
components of the dermis—the 
papillary layer and the reticular 
layer. Both are made of connective 
tissue with fibers of collagen 
extending from one to the other, 
making the border between the 
two somewhat indistinct. The 
dermal papillae extending into the 
epidermis belong to the papillary 
layer, whereas the dense collagen 
fiber bundles below belong to the 
reticular layer. LM x 10. (credit: 
modification of work by 
“kilbad”/Wikimedia Commons) 


Papillary Layer 


The papillary layer is made of loose, areolar connective tissue, which 
means the collagen and elastin fibers of this layer form a loose mesh. This 
superficial layer of the dermis projects into the stratum basale of the 
epidermis to form finger-like dermal papillae (see [link]). Within the 
papillary layer are fibroblasts, a small number of fat cells (adipocytes), and 
an abundance of small blood vessels. In addition, the papillary layer 
contains phagocytes, defensive cells that help fight bacteria or other 
infections that have breached the skin. This layer also contains lymphatic 
capillaries, nerve fibers, and touch receptors called the Meissner corpuscles. 


Reticular Layer 


Underlying the papillary layer is the much thicker reticular layer, 
composed of dense, irregular connective tissue. This layer is well 
vascularized and has a rich sensory and sympathetic nerve supply. The 
reticular layer appears reticulated (net-like) due to a tight meshwork of 
fibers. Elastin fibers provide some elasticity to the skin, enabling 
movement. Collagen fibers provide structure and tensile strength, with 
strands of collagen extending into both the papillary layer and the 
hypodermis. In addition, collagen binds water to keep the skin hydrated. 
Collagen injections and Retin-A creams help restore skin turgor by either 
introducing collagen externally or stimulating blood flow and repair of the 
dermis, respectively. 


Hypodermis 


The hypodermis (also called the subcutaneous layer or superficial fascia) is 
a layer directly below the dermis and serves to connect the skin to the 
underlying fascia (fibrous tissue) of the bones and muscles. It is not strictly 
a part of the skin, although the border between the hypodermis and dermis 
can be difficult to distinguish. The hypodermis consists of well- 
vascularized, loose, areolar connective tissue and adipose tissue, which 


functions as a mode of fat storage and provides insulation and cushioning 
for the integument. 


Note: 

Everyday Connection 

Lipid Storage 

The hypodermis is home to most of the fat that concerns people when they 
are trying to keep their weight under control. Adipose tissue present in the 
hypodermis consists of fat-storing cells called adipocytes. This stored fat 
can serve as an energy reserve, insulate the body to prevent heat loss, and 
act as a cushion to protect underlying structures from trauma. 

Where the fat is deposited and accumulates within the hypodermis depends 
on hormones (testosterone, estrogen, insulin, glucagon, leptin, and others), 
as well as genetic factors. Fat distribution changes as our bodies mature 
and age. Men tend to accumulate fat in different areas (neck, arms, lower 
back, and abdomen) than do women (breasts, hips, thighs, and buttocks). 
The body mass index (BMI) is often used as a measure of fat, although this 
measure is, in fact, derived from a mathematical formula that compares 
body weight (mass) to height. Therefore, its accuracy as a health indicator 
can be called into question in individuals who are extremely physically fit. 
In many animals, there is a pattern of storing excess calories as fat to be 
used in times when food is not readily available. In much of the developed 
world, insufficient exercise coupled with the ready availability and 
consumption of high-calorie foods have resulted in unwanted 
accumulations of adipose tissue in many people. Although periodic 
accumulation of excess fat may have provided an evolutionary advantage 
to our ancestors, who experienced unpredictable bouts of famine, it is now 
becoming chronic and considered a major health threat. Recent studies 
indicate that a distressing percentage of our population is overweight 
and/or clinically obese. Not only is this a problem for the individuals 
affected, but it also has a severe impact on our healthcare system. Changes 
in lifestyle, specifically in diet and exercise, are the best ways to control 
body fat accumulation, especially when it reaches levels that increase the 
risk of heart disease and diabetes. 


Pigmentation 


The color of skin is influenced by a number of pigments, including melanin, 
carotene, and hemoglobin. Recall that melanin is produced by cells called 
melanocytes, which are found scattered throughout the stratum basale of the 
epidermis. The melanin is transferred into the keratinocytes via a cellular 
vesicle called a melanosome ((link]). 

Skin Pigmentation 


Surface 


Upper 
keratinocytes 


Melanosomes 


Basal 
keratinocytes 


Melanocytes 


The relative coloration of the skin depends of the amount of 
melanin produced by melanocytes in the stratum basale and 
taken up by keratinocytes. 


Melanin occurs in two primary forms. Eumelanin exists as black and 
brown, whereas pheomelanin provides a red color. Dark-skinned 
individuals produce more melanin than those with pale skin. Exposure to 


the UV rays of the sun or a tanning salon causes melanin to be 
manufactured and built up in keratinocytes, as sun exposure stimulates 
keratinocytes to secrete chemicals that stimulate melanocytes. The 
accumulation of melanin in keratinocytes results in the darkening of the 
skin, or a tan. This increased melanin accumulation protects the DNA of 
epidermal cells from UV ray damage and the breakdown of folic acid, a 
nutrient necessary for our health and well-being. In contrast, too much 
melanin can interfere with the production of vitamin D, an important 
nutrient involved in calcium absorption. Thus, the amount of melanin 
present in our skin is dependent on a balance between available sunlight 
and folic acid destruction, and protection from UV radiation and vitamin D 
production. 


It requires about 10 days after initial sun exposure for melanin synthesis to 
peak, which is why pale-skinned individuals tend to suffer sunburns of the 
epidermis initially. Dark-skinned individuals can also get sunburns, but are 
more protected than are pale-skinned individuals. Melanosomes are 
temporary structures that are eventually destroyed by fusion with 
lysosomes; this fact, along with melanin-filled keratinocytes in the stratum 
corneum sloughing off, makes tanning impermanent. 


Too much sun exposure can eventually lead to wrinkling due to the 
destruction of the cellular structure of the skin, and in severe cases, can 
cause sufficient DNA damage to result in skin cancer. When there is an 
irregular accumulation of melanocytes in the skin, freckles appear. Moles 
are larger masses of melanocytes, and although most are benign, they 
should be monitored for changes that might indicate the presence of cancer 
({link]). 

Moles 


Moles range from benign accumulations 
of melanocytes to melanomas. These 
structures populate the landscape of our 
skin. (credit: the National Cancer 
Institute) 


Note: 

Disorders of the... 

Integumentary System 

The first thing a clinician sees is the skin, and so the examination of the 
skin should be part of any thorough physical examination. Most skin 
disorders are relatively benign, but a few, including melanomas, can be 
fatal if untreated. A couple of the more noticeable disorders, albinism and 
vitiligo, affect the appearance of the skin and its accessory organs. 
Although neither is fatal, it would be hard to claim that they are benign, at 
least to the individuals so afflicted. 


Albinism is a genetic disorder that affects (completely or partially) the 
coloring of skin, hair, and eyes. The defect is primarily due to the inability 
of melanocytes to produce melanin. Individuals with albinism tend to 
appear white or very pale due to the lack of melanin in their skin and hair. 
Recall that melanin helps protect the skin from the harmful effects of UV 
radiation. Individuals with albinism tend to need more protection from UV 
radiation, as they are more prone to sunburns and skin cancer. They also 
tend to be more sensitive to light and have vision problems due to the lack 
of pigmentation on the retinal wall. Treatment of this disorder usually 
involves addressing the symptoms, such as limiting UV light exposure to 
the skin and eyes. In vitiligo, the melanocytes in certain areas lose their 
ability to produce melanin, possibly due to an autoimmune reaction. This 
leads to a loss of color in patches ({link]). Neither albinism nor vitiligo 
directly affects the lifespan of an individual. 

Vitiligo 


Individuals with 
vitiligo experience 
depigmentation that 
results in lighter 
colored patches of skin. 
The condition is 
especially noticeable 


on darker skin. (credit: 
Klaus D. Peter) 


Other changes in the appearance of skin coloration can be indicative of 
diseases associated with other body systems. Liver disease or liver cancer 
can cause the accumulation of bile and the yellow pigment bilirubin, 
leading to the skin appearing yellow or jaundiced (jaune is the French 
word for “yellow”). Tumors of the pituitary gland can result in the 
secretion of large amounts of melanocyte-stimulating hormone (MSH), 
which results in a darkening of the skin. Similarly, Addison’s disease can 
stimulate the release of excess amounts of adrenocorticotropic hormone 
(ACTH), which can give the skin a deep bronze color. A sudden drop in 
oxygenation can affect skin color, causing the skin to initially turn ashen 
(white). With a prolonged reduction in oxygen levels, dark red 
deoxyhemoglobin becomes dominant in the blood, making the skin appear 
blue, a condition referred to as cyanosis (kyanos is the Greek word for 
“blue”). This happens when the oxygen supply is restricted, as when 
someone is experiencing difficulty in breathing because of asthma or a 
heart attack. However, in these cases the effect on skin color has nothing 
do with the skin’s pigmentation. 


mess’ OPENStAX COLLEGE” 
seh : T 


This ABC video follows the story of a pair of fraternal African-American 
twins, one of whom is albino. Watch this video to learn about the 
challenges these children and their family face. Which ethnicities do you 
think are exempt from the possibility of albinism? 


Chapter Review 


The skin is composed of two major layers: a superficial epidermis and a 
deeper dermis. The epidermis consists of several layers beginning with the 
innermost (deepest) stratum basale (germinatum), followed by the stratum 
spinosum, stratum granulosum, stratum lucidum (when present), and ending 
with the outermost layer, the stratum corneum. The topmost layer, the 
stratum corneum, consists of dead cells that shed periodically and is 
progressively replaced by cells formed from the basal layer. The stratum 
basale also contains melanocytes, cells that produce melanin, the pigment 
primarily responsible for giving skin its color. Melanin is transferred to 
keratinocytes in the stratum spinosum to protect cells from UV rays. 


The dermis connects the epidermis to the hypodermis, and provides strength 
and elasticity due to the presence of collagen and elastin fibers. It has only 
two layers: the papillary layer with papillae that extend into the epidermis 
and the lower, reticular layer composed of loose connective tissue. The 
hypodermis, deep to the dermis of skin, is the connective tissue that 
connects the dermis to underlying structures; it also harbors adipose tissue 
for fat storage and protection. 


Interactive Link Questions 


Exercise: 
Problem: 
The skin consists of two layers and a closely associated layer. View 


this animation to learn more about layers of the skin. What are the 
basic functions of each of these layers? 


Solution: 


The epidermis provides protection, the dermis provides support and 
flexibility, and the hypodermis (fat layer) provides insulation and 
padding. 


Exercise: 


Problem: 


[link] If you zoom on the cells at the outermost layer of this section of 
skin, what do you notice about the cells? 


Solution: 


[link] These cells do not have nuclei, so you can deduce that they are 
dead. They appear to be sloughing off. 

Exercise: 
Problem: 


[link] If you zoom on the cells of the stratum spinosum, what is 
distinctive about them? 


Solution: 


[link] These cells have desmosomes, which give the cells their spiny 
appearance. 


Exercise: 
Problem: 
This ABC video follows the story of a pair of fraternal African- 
American twins, one of whom is albino. Watch this video to learn 
about the challenges these children and their family face. Which 
ethnicities do you think are exempt from the possibility of albinism? 


Solution: 


There are none. 


Review Questions 


Exercise: 


Problem: 


The papillary layer of the dermis is most closely associated with which 
layer of the epidermis? 


a. stratum spinosum 
b. stratum corneum 

c. stratum granulosum 
d. stratum basale 


Solution: 


D 


Exercise: 


Problem:Langerhans cells are commonly found in the 


a. stratum spinosum 
b. stratum corneum 

c. stratum granulosum 
d. stratum basale 


Solution: 


A 
Exercise: 
Problem: 


The papillary and reticular layers of the dermis are composed mainly 
of 


a. melanocytes 
b. keratinocytes 
c. connective tissue 


d. adipose tissue 


Solution: 
C 
Exercise: 
Problem: Collagen lends to the skin. 
a. elasticity 
b. structure 


c. color 
d. UV protection 


Solution: 


B 


Exercise: 


Problem: Which of the following is not a function of the hypodermis? 
a. protects underlying organs 
b. helps maintain body temperature 
c. source of blood vessels in the epidermis 
d. a site to long-term energy storage 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: 


What determines the color of skin, and what is the process that darkens 
skin when it is exposed to UV light? 


Solution: 


The pigment melanin, produced by melanocytes, is primarily 
responsible for skin color. Melanin comes in different shades of brown 
and black. Individuals with darker skin have darker, more abundant 
melanin, whereas fair-skinned individuals have a lighter shade of skin 
and less melanin. Exposure to UV irradiation stimulates the 
melanocytes to produce and secrete more melanin. 


Exercise: 


Problem: 


Cells of the epidermis derive from stem cells of the stratum basale. 
Describe how the cells change as they become integrated into the 
different layers of the epidermis. 


Solution: 


As the cells move into the stratum spinosum, they begin the synthesis 
of keratin and extend cell processes, desmosomes, which link the cells. 
As the stratum basale continues to produce new cells, the keratinocytes 
of the stratum spinosum are pushed into the stratum granulosum. The 
cells become flatter, their cell membranes thicken, and they generate 
large amounts of the proteins keratin and keratohyalin. The nuclei and 
other cell organelles disintegrate as the cells die, leaving behind the 
keratin, keratohyalin, and cell membranes that form the stratum 
lucidum and the stratum corneum. The keratinocytes in these layers are 
mostly dead and flattened. Cells in the stratum corneum are 
periodically shed. 


Glossary 


albinism 
genetic disorder that affects the skin, in which there is no melanin 
production 


basal cell 
type of stem cell found in the stratum basale and in the hair matrix that 
continually undergoes cell division, producing the keratinocytes of the 
epidermis 


dermal papilla 
(plural = dermal papillae) extension of the papillary layer of the dermis 
that increases surface contact between the epidermis and dermis 


dermis 
layer of skin between the epidermis and hypodermis, composed mainly 
of connective tissue and containing blood vessels, hair follicles, sweat 
glands, and other structures 


desmosome 
structure that forms an impermeable junction between cells 


elastin fibers 
fibers made of the protein elastin that increase the elasticity of the 
dermis 


eleiden 
clear protein-bound lipid found in the stratum lucidum that is derived 
from keratohyalin and helps to prevent water loss 


epidermis 
outermost tissue layer of the skin 


hypodermis 
connective tissue connecting the integument to the underlying bone 
and muscle 


integumentary system 
skin and its accessory structures 


keratin 
type of structural protein that gives skin, hair, and nails its hard, water- 
resistant properties 


keratinocyte 
cell that produces keratin and is the most predominant type of cell 
found in the epidermis 


keratohyalin 
granulated protein found in the stratum granulosum 


Langerhans cell 
specialized dendritic cell found in the stratum spinosum that functions 
as a macrophage 


melanin 
pigment that determines the color of hair and skin 


melanocyte 
cell found in the stratum basale of the epidermis that produces the 
pigment melanin 


melanosome 
intercellular vesicle that transfers melanin from melanocytes into 
keratinocytes of the epidermis 


Merkel cell 
receptor cell in the stratum basale of the epidermis that responds to the 
sense of touch 


papillary layer 
superficial layer of the dermis, made of loose, areolar connective tissue 


reticular layer 
deeper layer of the dermis; it has a reticulated appearance due to the 
presence of abundant collagen and elastin fibers 


stratum basale 


deepest layer of the epidermis, made of epidermal stem cells 


stratum corneum 
most superficial layer of the epidermis 


stratum granulosum 
layer of the epidermis superficial to the stratum spinosum 


stratum lucidum 
layer of the epidermis between the stratum granulosum and stratum 
corneum, found only in thick skin covering the palms, soles of the feet, 
and digits 


stratum spinosum 
layer of the epidermis superficial to the stratum basale, characterized 
by the presence of desmosomes 


vitiligo 
skin condition in which melanocytes in certain areas lose the ability to 
produce melanin, possibly due an autoimmune reaction that leads to 
loss of color in patches 


Accessory Structures of the Skin 
By the end of this section, you will be able to: 


e Identify the accessory structures of the skin 

e Describe the structure and function of hair and nails 

e Describe the structure and function of sweat glands and sebaceous 
glands 


Accessory structures of the skin include hair, nails, sweat glands, and 
sebaceous glands. These structures embryologically originate from the 
epidermis and can extend down through the dermis into the hypodermis. 


Hair 


Hair is a keratinous filament growing out of the epidermis. It is primarily 
made of dead, keratinized cells. Strands of hair originate in an epidermal 
penetration of the dermis called the hair follicle. The hair shaft is the part 
of the hair not anchored to the follicle, and much of this is exposed at the 
skin’s surface. The rest of the hair, which is anchored in the follicle, lies 
below the surface of the skin and is referred to as the hair root. The hair 
root ends deep in the dermis at the hair bulb, and includes a layer of 
mitotically active basal cells called the hair matrix. The hair bulb 
surrounds the hair papilla, which is made of connective tissue and contains 
blood capillaries and nerve endings from the dermis ([link]). 

Hair 


Medulla 
Cortex 


Cuticle 


Sebaceous 
gland 


Inner root 
sheath 


Outer root 
sheath 


Hair matrix 


Hair 
papilla 


Hair follicles originate in the 
epidermis and have many 
different parts. 


Just as the basal layer of the epidermis forms the layers of epidermis that 
get pushed to the surface as the dead skin on the surface sheds, the basal 
cells of the hair bulb divide and push cells outward in the hair root and shaft 
as the hair grows. The medulla forms the central core of the hair, which is 
surrounded by the cortex, a layer of compressed, keratinized cells that is 
covered by an outer layer of very hard, keratinized cells known as the 
cuticle. These layers are depicted in a longitudinal cross-section of the hair 
follicle ([link]), although not all hair has a medullary layer. Hair texture 
(straight, curly) is determined by the shape and structure of the cortex, and 
to the extent that it is present, the medulla. The shape and structure of these 
layers are, in turn, determined by the shape of the hair follicle. Hair growth 
begins with the production of keratinocytes by the basal cells of the hair 
bulb. As new cells are deposited at the hair bulb, the hair shaft is pushed 
through the follicle toward the surface. Keratinization is completed as the 


cells are pushed to the skin surface to form the shaft of hair that is 
externally visible. The external hair is completely dead and composed 
entirely of keratin. For this reason, our hair does not have sensation. 
Furthermore, you can cut your hair or shave without damaging the hair 
structure because the cut is superficial. Most chemical hair removers also 
act superficially; however, electrolysis and yanking both attempt to destroy 
the hair bulb so hair cannot grow. 

Hair Follicle 


The slide shows a cross-section of a hair 
follicle. Basal cells of the hair matrix in 
the center differentiate into cells of the 
inner root sheath. Basal cells at the base 

of the hair root form the outer root sheath. 

LM x 4. (credit: modification of work by 

“kilbad”/Wikimedia Commons) 


The wall of the hair follicle is made of three concentric layers of cells. The 
cells of the internal root sheath surround the root of the growing hair and 
extend just up to the hair shaft. They are derived from the basal cells of the 
hair matrix. The external root sheath, which is an extension of the 
epidermis, encloses the hair root. It is made of basal cells at the base of the 
hair root and tends to be more keratinous in the upper regions. The glassy 


membrane is a thick, clear connective tissue sheath covering the hair root, 
connecting it to the tissue of the dermis. 


Note: 


Dasara 


— 
meee, OPENStAX COLLEGE 
—— . 


The hair follicle is made of multiple layers of cells that form from basal 
cells in the hair matrix and the hair root. Cells of the hair matrix divide and 
differentiate to form the layers of the hair. Watch this video to learn more 
about hair follicles. 


Hair serves a variety of functions, including protection, sensory input, 
thermoregulation, and communication. For example, hair on the head 
protects the skull from the sun. The hair in the nose and ears, and around 
the eyes (eyelashes) defends the body by trapping and excluding dust 
particles that may contain allergens and microbes. Hair of the eyebrows 
prevents sweat and other particles from dripping into and bothering the 
eyes. Hair also has a sensory function due to sensory innervation by a hair 
root plexus surrounding the base of each hair follicle. Hair is extremely 
sensitive to air movement or other disturbances in the environment, much 
more so than the skin surface. This feature is also useful for the detection of 
the presence of insects or other potentially damaging substances on the skin 
surface. Each hair root is connected to a smooth muscle called the arrector 
pili that contracts in response to nerve signals from the sympathetic nervous 
system, making the external hair shaft “stand up.” The primary purpose for 
this is to trap a layer of air to add insulation. This is visible in humans as 
goose bumps and even more obvious in animals, such as when a frightened 
cat raises its fur. Of course, this is much more obvious in organisms with a 
heavier coat than most humans, such as dogs and cats. 


Hair Growth 


Hair grows and is eventually shed and replaced by new hair. This occurs in 
three phases. The first is the anagen phase, during which cells divide 
rapidly at the root of the hair, pushing the hair shaft up and out. The length 
of this phase is measured in years, typically from 2 to 7 years. The catagen 
phase lasts only 2 to 3 weeks, and marks a transition from the hair follicle’s 
active growth. Finally, during the telogen phase, the hair follicle is at rest 
and no new growth occurs. At the end of this phase, which lasts about 2 to 4 
months, another anagen phase begins. The basal cells in the hair matrix then 
produce a new hair follicle, which pushes the old hair out as the growth 
cycle repeats itself. Hair typically grows at the rate of 0.3 mm per day 
during the anagen phase. On average, 50 hairs are lost and replaced per day. 
Hair loss occurs if there is more hair shed than what is replaced and can 
happen due to hormonal or dietary changes. Hair loss can also result from 
the aging process, or the influence of hormones. 


Hair Color 


Similar to the skin, hair gets its color from the pigment melanin, produced 
by melanocytes in the hair papilla. Different hair color results from 
differences in the type of melanin, which is genetically determined. As a 
person ages, the melanin production decreases, and hair tends to lose its 
color and becomes gray and/or white. 


Nails 


The nail bed is a specialized structure of the epidermis that is found at the 
tips of our fingers and toes. The nail body is formed on the nail bed, and 
protects the tips of our fingers and toes as they are the farthest extremities 
and the parts of the body that experience the maximum mechanical stress 
({link]). In addition, the nail body forms a back-support for picking up 

small objects with the fingers. The nail body is composed of densely packed 
dead keratinocytes. The epidermis in this part of the body has evolved a 
specialized structure upon which nails can form. The nail body forms at the 


nail root, which has a matrix of proliferating cells from the stratum basale 
that enables the nail to grow continuously. The lateral nail fold overlaps the 
nail on the sides, helping to anchor the nail body. The nail fold that meets 
the proximal end of the nail body forms the nail cuticle, also called the 
eponychium. The nail bed is rich in blood vessels, making it appear pink, 
except at the base, where a thick layer of epithelium over the nail matrix 
forms a crescent-shaped region called the lunula (the “little moon”). The 
area beneath the free edge of the nail, furthest from the cuticle, is called the 
hyponychium. It consists of a thickened layer of stratum corneum. 

Nails 


Free edge Eponychium 
a ee Proximal nail fold 


. Lunula 
Nail / 


Nail body 
Lateral nail fold 


Lunula 
Eponychium 
Proximal nail fold 


Epidermis Dermis Phalanx Hyponychium 


The nail is an accessory structure of the integumentary 
system. 


Note: 


— e 
meee <OPENStAX COLLEGE” 
— 


wy 


oe atten 


Nails are accessory structures of the integumentary system. Visit this link 
to learn more about the origin and growth of fingernails. 


Sweat Glands 


When the body becomes warm, sudoriferous glands produce sweat to cool 
the body. Sweat glands develop from epidermal projections into the dermis 
and are classified as merocrine glands; that is, the secretions are excreted by 
exocytosis through a duct without affecting the cells of the gland. There are 
two types of sweat glands, each secreting slightly different products. 


An eccrine sweat gland is type of gland that produces a hypotonic sweat 
for thermoregulation. These glands are found all over the skin’s surface, but 
are especially abundant on the palms of the hand, the soles of the feet, and 
the forehead ([link]). They are coiled glands lying deep in the dermis, with 
the duct rising up to a pore on the skin surface, where the sweat is released. 
This type of sweat, released by exocytosis, is hypotonic and composed 
mostly of water, with some salt, antibodies, traces of metabolic waste, and 
dermicidin, an antimicrobial peptide. Eccrine glands are a primary 
component of thermoregulation in humans and thus help to maintain 
homeostasis. 

Eccrine Gland 


Skin surface 


Eccrine sweat 
gland 


Eccrine glands are coiled glands in the 
dermis that release sweat that is 


mostly water. 


An apocrine sweat gland is usually associated with hair follicles in densely 
hairy areas, such as armpits and genital regions. Apocrine sweat glands are 
larger than eccrine sweat glands and lie deeper in the dermis, sometimes 
even reaching the hypodermis, with the duct normally emptying into the 
hair follicle. In addition to water and salts, apocrine sweat includes organic 
compounds that make the sweat thicker and subject to bacterial 
decomposition and subsequent smell. The release of this sweat is under 
both nervous and hormonal control, and plays a role in the poorly 
understood human pheromone response. Most commercial antiperspirants 
use an aluminum-based compound as their primary active ingredient to stop 
sweat. When the antiperspirant enters the sweat gland duct, the aluminum- 
based compounds precipitate due to a change in pH and form a physical 
block in the duct, which prevents sweat from coming out of the pore. 


— aa COLLEGE” 


Sweating regulates body temperature. The composition of the sweat 
determines whether body odor is a byproduct of sweating. Visit this link to 
learn more about sweating and body odor. 


Sebaceous Glands 


A sebaceous gland is a type of oil gland that is found all over the body and 
helps to lubricate and waterproof the skin and hair. Most sebaceous glands 


are associated with hair follicles. They generate and excrete sebum, a 
mixture of lipids, onto the skin surface, thereby naturally lubricating the dry 
and dead layer of keratinized cells of the stratum corneum, keeping it 
pliable. The fatty acids of sebum also have antibacterial properties, and 
prevent water loss from the skin in low-humidity environments. The 
secretion of sebum is stimulated by hormones, many of which do not 
become active until puberty. Thus, sebaceous glands are relatively inactive 
during childhood. 


Chapter Review 


Accessory structures of the skin include hair, nails, sweat glands, and 
sebaceous glands. Hair is made of dead keratinized cells, and gets its color 
from melanin pigments. Nails, also made of dead keratinized cells, protect 
the extremities of our fingers and toes from mechanical damage. Sweat 
glands and sebaceous glands produce sweat and sebum, respectively. Each 
of these fluids has a role to play in maintaining homeostasis. Sweat cools 
the body surface when it gets overheated and helps excrete small amounts 
of metabolic waste. Sebum acts as a natural moisturizer and keeps the dead, 
flaky, outer keratin layer healthy. 


Review Questions 


Exercise: 
Problem: 


In response to stimuli from the sympathetic nervous system, the 
arrector pili 


a. are glands on the skin surface 

b. can lead to excessive sweating 
c. are responsible for goose bumps 
d. secrete sebum 


Solution: 


C 


Exercise: 


Problem:The hair matrix contains 


a. the hair follicle 

b. the hair shaft 

c. the glassy membrane 
d. a layer of basal cells 


Solution: 


D 


Exercise: 


Problem:Eccrine sweat glands 


a. are present on hair 

b. are present in the skin throughout the body and produce watery 
Sweat 

c. produce sebum 

d. act as a moisturizer 


Solution: 


B 


Exercise: 


Problem: Sebaceous glands 


a. are a type of sweat gland 

b. are associated with hair follicles 

c. may function in response to touch 

d. release a watery solution of salt and metabolic waste 


Solution: 


B 
Exercise: 


Problem: 


Similar to the hair, nails grow continuously throughout our lives. 
Which of the following is furthest from the nail growth center? 


a. nail bed 

b. hyponychium 
c. nail root 

d. eponychium 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


Explain the differences between eccrine and apocrine sweat glands. 


Solution: 


Eccrine sweat glands are all over the body, especially the forehead and 
palms of the hand. They release a watery sweat, mixed with some 
metabolic waste and antibodies. Apocrine glands are associated with 
hair follicles. They are larger than eccrine sweat glands and lie deeper 
in the dermis, sometimes even reaching the hypodermis. They release 


a thicker sweat that is often decomposed by bacteria on the skin, 
resulting in an unpleasant odor. 


Exercise: 


Problem: Describe the structure and composition of nails. 


Solution: 


Nails are composed of densely packed dead keratinocytes. They 
protect the fingers and toes from mechanical stress. The nail body is 
formed on the nail bed, which is at the nail root. Nail folds, folds of 
skin that overlap the nail on its side, secure the nail to the body. The 
crescent-shaped region at the base of the nail is the lunula. 


Glossary 


anagen 
active phase of the hair growth cycle 


apocrine sweat gland 
type of sweat gland that is associated with hair follicles in the armpits 
and genital regions 


arrector pili 
smooth muscle that is activated in response to external stimuli that pull 
on hair follicles and make the hair “stand up” 


catagen 
transitional phase marking the end of the anagen phase of the hair 
growth cycle 


cortex 
in hair, the second or middle layer of keratinocytes originating from 
the hair matrix, as seen in a cross-section of the hair bulb 


cuticle 


in hair, the outermost layer of keratinocytes originating from the hair 
matrix, as seen in a cross-section of the hair bulb 


eccrine sweat gland 
type of sweat gland that is common throughout the skin surface; it 
produces a hypotonic sweat for thermoregulation 


eponychium 
nail fold that meets the proximal end of the nail body, also called the 
cuticle 


external root sheath 
outer layer of the hair follicle that is an extension of the epidermis, 
which encloses the hair root 


glassy membrane 
layer of connective tissue that surrounds the base of the hair follicle, 
connecting it to the dermis 


hair 
keratinous filament growing out of the epidermis 


hair bulb 
structure at the base of the hair root that surrounds the dermal papilla 


hair follicle 
cavity or sac from which hair originates 


hair matrix 
layer of basal cells from which a strand of hair grows 


hair papilla 
mass of connective tissue, blood capillaries, and nerve endings at the 


base of the hair follicle 


hair root 
part of hair that is below the epidermis anchored to the follicle 


hair shaft 


part of hair that is above the epidermis but is not anchored to the 
follicle 


hyponychium 
thickened layer of stratum corneum that lies below the free edge of the 
nail 


internal root sheath 
innermost layer of keratinocytes in the hair follicle that surround the 
hair root up to the hair shaft 


lunula 
basal part of the nail body that consists of a crescent-shaped layer of 
thick epithelium 


medulla 
in hair, the innermost layer of keratinocytes originating from the hair 
matrix 


nail bed 
layer of epidermis upon which the nail body forms 


nail body 
main keratinous plate that forms the nail 


nail cuticle 
fold of epithelium that extends over the nail bed, also called the 
eponychium 


nail fold 
fold of epithelium at that extend over the sides of the nail body, 
holding it in place 


nail root 
part of the nail that is lodged deep in the epidermis from which the nail 
gTOWS 


sebaceous gland 


type of oil gland found in the dermis all over the body and helps to 
lubricate and waterproof the skin and hair by secreting sebum 


sebum 
oily substance that is composed of a mixture of lipids that lubricates 
the skin and hair 


sudoriferous gland 
sweat gland 


telogen 
resting phase of the hair growth cycle initiated with catagen and 
terminated by the beginning of a new anagen phase of hair growth 


Diseases, Disorders, and Injuries of the Integumentary System 
By the end of this section, you will be able to: 


e Describe several different diseases and disorders of the skin 
e Describe the effect of injury to the skin and the process of healing 


The integumentary system is susceptible to a variety of diseases, disorders, 
and injuries. These range from annoying but relatively benign bacterial or 
fungal infections that are categorized as disorders, to skin cancer and severe 
burns, which can be fatal. In this section, you will learn several of the most 
common skin conditions. 


Diseases 


One of the most talked about diseases is skin cancer. Cancer is a broad term 
that describes diseases caused by abnormal cells in the body dividing 
uncontrollably. Most cancers are identified by the organ or tissue in which 
the cancer originates. One common form of cancer is skin cancer. The Skin 
Cancer Foundation reports that one in five Americans will experience some 
type of skin cancer in their lifetime. The degradation of the ozone layer in 
the atmosphere and the resulting increase in exposure to UV radiation has 
contributed to its rise. Overexposure to UV radiation damages DNA, which 
can lead to the formation of cancerous lesions. Although melanin offers 
some protection against DNA damage from the sun, often it is not enough. 
The fact that cancers can also occur on areas of the body that are normally 
not exposed to UV radiation suggests that there are additional factors that 
can lead to cancerous lesions. 


In general, cancers result from an accumulation of DNA mutations. These 
mutations can result in cell populations that do not die when they should 
and uncontrolled cell proliferation that leads to tumors. Although many 
tumors are benign (harmless), some produce cells that can mobilize and 
establish tumors in other organs of the body; this process is referred to as 
metastasis. Cancers are characterized by their ability to metastasize. 


Basal Cell Carcinoma 


Basal cell carcinoma is a form of cancer that affects the mitotically active 
stem cells in the stratum basale of the epidermis. It is the most common of 
all cancers that occur in the United States and is frequently found on the 
head, neck, arms, and back, which are areas that are most susceptible to 
long-term sun exposure. Although UV rays are the main culprit, exposure to 
other agents, such as radiation and arsenic, can also lead to this type of 
cancer. Wounds on the skin due to open sores, tattoos, burns, etc. may be 
predisposing factors as well. Basal cell carcinomas start in the stratum 
basale and usually spread along this boundary. At some point, they begin to 
grow toward the surface and become an uneven patch, bump, growth, or 
scar on the skin surface ([link]). Like most cancers, basal cell carcinomas 
respond best to treatment when caught early. Treatment options include 
surgery, freezing (cryosurgery), and topical ointments (Mayo Clinic 2012). 
Basal Cell Carcinoma 


Basal cell carcinoma 


can take several 
different forms. Similar 
to other forms of skin 
cancer, it is readily 
cured if caught early 
and treated. (credit: 
John Hendrix, MD) 


Squamous Cell Carcinoma 


Squamous cell carcinoma is a cancer that affects the keratinocytes of the 
stratum spinosum and presents as lesions commonly found on the scalp, 
ears, and hands ([link]). It is the second most common skin cancer. The 
American Cancer Society reports that two of 10 skin cancers are squamous 
cell carcinomas, and it is more aggressive than basal cell carcinoma. If not 
removed, these carcinomas can metastasize. Surgery and radiation are used 
to cure squamous cell carcinoma. 

Squamous Cell Carcinoma 


Squamous cell carcinoma 
presents here as a lesion on 
an individual’s nose. (credit: 

the National Cancer Institute) 


Melanoma 


A melanoma is a cancer characterized by the uncontrolled growth of 
melanocytes, the pigment-producing cells in the epidermis. Typically, a 
melanoma develops from a mole. It is the most fatal of all skin cancers, as it 
is highly metastatic and can be difficult to detect before it has spread to 
other organs. Melanomas usually appear as asymmetrical brown and black 
patches with uneven borders and a raised surface ([{link]). Treatment 
typically involves surgical excision and immunotherapy. 

Melanoma 


Melanomas typically 
present as large brown 
or black patches with 
uneven borders and a 
raised surface. (credit: 
the National Cancer 


Institute) 


Doctors often give their patients the following ABCDE mnemonic to help 
with the diagnosis of early-stage melanoma. If you observe a mole on your 
body displaying these signs, consult a doctor. 


e Asymmetry — the two sides are not symmetrical 

¢ Borders — the edges are irregular in shape 

e Color — the color is varied shades of brown or black 
e Diameter — it is larger than 6 mm (0.24 in) 

e Evolving — its shape has changed 


Some specialists cite the following additional signs for the most serious 
form, nodular melanoma: 


e Elevated — it is raised on the skin surface 
e Firm — it feels hard to the touch 
¢ Growing — it is getting larger 


Skin Disorders 


Two common skin disorders are eczema and acne. Eczema is an 
inflammatory condition and occurs in individuals of all ages. Acne involves 
the clogging of pores, which can lead to infection and inflammation, and is 
often seen in adolescents. Other disorders, not discussed here, include 
seborrheic dermatitis (on the scalp), psoriasis, cold sores, impetigo, scabies, 
hives, and warts. 


Eczema 


Eczema is an allergic reaction that manifests as dry, itchy patches of skin 
that resemble rashes ([link]). It may be accompanied by swelling of the 
skin, flaking, and in severe cases, bleeding. Many who suffer from eczema 
have antibodies against dust mites in their blood, but the link between 
eczema and allergy to dust mites has not been proven. Symptoms are 
usually managed with moisturizers, corticosteroid creams, and 
immunosuppressants. 

Eczema 


Eczema is a common skin 
disorder that presents as a 
red, flaky rash. (credit: 
“Jambula”/Wikimedia 
Commons) 


Acne 


Acne is a skin disturbance that typically occurs on areas of the skin that are 
rich in sebaceous glands (face and back). It is most common along with the 
onset of puberty due to associated hormonal changes, but can also occur in 
infants and continue into adulthood. Hormones, such as androgens, 
stimulate the release of sebum. An overproduction and accumulation of 
sebum along with keratin can block hair follicles. This plug is initially 
white. The sebum, when oxidized by exposure to air, turns black. Acne 
results from infection by acne-causing bacteria (Propionibacterium and 
Staphylococcus), which can lead to redness and potential scarring due to the 
natural wound healing process ((Link]). 


Acne 
Epidermis Plugged follicle Mild inflammation Marked inflammation 


Sebaceous 


Accumulation of shed Bacteria proliferate 
keratin and sebum 


Acne is a result of over-productive sebaceous glands, 


which leads to formation of blackheads and 
inflammation of the skin. 


Note: 


Career Connection 

Dermatologist 

Have you ever had a skin rash that did not respond to over-the-counter 
creams, or a mole that you were concerned about? Dermatologists help 
patients with these types of problems and more, on a daily basis. 
Dermatologists are medical doctors who specialize in diagnosing and 
treating skin disorders. Like all medical doctors, dermatologists earn a 
medical degree and then complete several years of residency training. In 
addition, dermatologists may then participate in a dermatology fellowship 
or complete additional, specialized training in a dermatology practice. If 
practicing in the United States, dermatologists must pass the United States 
Medical Licensing Exam (USMLE), become licensed in their state of 
practice, and be certified by the American Board of Dermatology. 

Most dermatologists work in a medical office or private-practice setting. 
They diagnose skin conditions and rashes, prescribe oral and topical 
medications to treat skin conditions, and may perform simple procedures, 
such as mole or wart removal. In addition, they may refer patients to an 
oncologist if skin cancer that has metastasized is suspected. Recently, 
cosmetic procedures have also become a prominent part of dermatology. 
Botox injections, laser treatments, and collagen and dermal filler injections 
are popular among patients, hoping to reduce the appearance of skin aging. 
Dermatology is a competitive specialty in medicine. Limited openings in 
dermatology residency programs mean that many medical students 
compete for a few select spots. Dermatology is an appealing specialty to 
many prospective doctors, because unlike emergency room physicians or 
surgeons, dermatologists generally do not have to work excessive hours or 
be “on-call” weekends and holidays. Moreover, the popularity of cosmetic 
dermatology has made it a growing field with many lucrative 
opportunities. It is not unusual for dermatology clinics to market 
themselves exclusively as cosmetic dermatology centers, and for 
dermatologists to specialize exclusively in these procedures. 

Consider visiting a dermatologist to talk about why he or she entered the 
field and what the field of dermatology is like. Visit this site for additional 
information. 


Injuries 


Because the skin is the part of our bodies that meets the world most directly, 
it is especially vulnerable to injury. Injuries include burns and wounds, as 
well as scars and calluses. They can be caused by sharp objects, heat, or 
excessive pressure or friction to the skin. 


Skin injuries set off a healing process that occurs in several overlapping 
stages. The first step to repairing damaged skin is the formation of a blood 
clot that helps stop the flow of blood and scabs over with time. Many 
different types of cells are involved in wound repair, especially if the 
surface area that needs repair is extensive. Before the basal stem cells of the 
stratum basale can recreate the epidermis, fibroblasts mobilize and divide 
rapidly to repair the damaged tissue by collagen deposition, forming 
granulation tissue. Blood capillaries follow the fibroblasts and help increase 
blood circulation and oxygen supply to the area. Immune cells, such as 
macrophages, roam the area and engulf any foreign matter to reduce the 
chance of infection. 


Burns 


A burn results when the skin is damaged by intense heat, radiation, 
electricity, or chemicals. The damage results in the death of skin cells, 
which can lead to a massive loss of fluid. Dehydration, electrolyte 
imbalance, and renal and circulatory failure follow, which can be fatal. Burn 
patients are treated with intravenous fluids to offset dehydration, as well as 
intravenous nutrients that enable the body to repair tissues and replace lost 
proteins. Another serious threat to the lives of burn patients is infection. 
Burned skin is extremely susceptible to bacteria and other pathogens, due to 
the loss of protection by intact layers of skin. 


Burns are sometimes measured in terms of the size of the total surface area 
affected. This is referred to as the “rule of nines,” which associates specific 
anatomical areas with a percentage that is a factor of nine ([link]). Burns are 
also classified by the degree of their severity. A first-degree burn is a 
superficial burn that affects only the epidermis. Although the skin may be 


painful and swollen, these burns typically heal on their own within a few 
days. Mild sunburn fits into the category of a first-degree burn. A second- 
degree burn goes deeper and affects both the epidermis and a portion of 
the dermis. These burns result in swelling and a painful blistering of the 
skin. It is important to keep the burn site clean and sterile to prevent 
infection. If this is done, the burn will heal within several weeks. A third- 
degree burn fully extends into the epidermis and dermis, destroying the 
tissue and affecting the nerve endings and sensory function. These are 
serious burns that may appear white, red, or black; they require medical 
attention and will heal slowly without it. A fourth-degree burn is even 
more severe, affecting the underlying muscle and bone. Oddly, third and 
fourth-degree burns are usually not as painful because the nerve endings 
themselves are damaged. Full-thickness burns cannot be repaired by the 
body, because the local tissues used for repair are damaged and require 
excision (debridement), or amputation in severe cases, followed by grafting 
of the skin from an unaffected part of the body, or from skin grown in tissue 
culture for grafting purposes. 

Calculating the Size of a Burn 


Head and 
neck 9% 


Upper limbs 
9% each 


Trunk 36% 


Genitalia 1% 


Lower limbs 
18% each 


The size of a burn will guide decisions 


made about the need for specialized 

treatment. Specific parts of the body 

are associated with a percentage of 
body area. 


me oS 
= openstax COLLEGE” 
r 4 
abe 


Skin grafts are required when the damage from trauma or infection cannot 
be closed with sutures or staples. Watch this video to learn more about skin 


grafting procedures. 


Scars and Keloids 


Most cuts or wounds, with the exception of ones that only scratch the 
surface (the epidermis), lead to scar formation. A scar is collagen-rich skin 
formed after the process of wound healing that differs from normal skin. 
Scarring occurs in cases in which there is repair of skin damage, but the 
skin fails to regenerate the original skin structure. Fibroblasts generate scar 
tissue in the form of collagen, and the bulk of repair is due to the basket- 
weave pattern generated by collagen fibers and does not result in 
regeneration of the typical cellular structure of skin. Instead, the tissue is 
fibrous in nature and does not allow for the regeneration of accessory 
structures, such as hair follicles, sweat glands, or sebaceous glands. 


Sometimes, there is an overproduction of scar tissue, because the process of 
collagen formation does not stop when the wound is healed; this results in 


the formation of a raised or hypertrophic scar called a keloid. In contrast, 
scars that result from acne and chickenpox have a sunken appearance and 
are called atrophic scars. 


Scarring of skin after wound healing is a natural process and does not need 
to be treated further. Application of mineral oil and lotions may reduce the 
formation of scar tissue. However, modern cosmetic procedures, such as 
dermabrasion, laser treatments, and filler injections have been invented as 
remedies for severe scarring. All of these procedures try to reorganize the 
structure of the epidermis and underlying collagen tissue to make it look 
more natural. 


Bedsores and Stretch Marks 


Skin and its underlying tissue can be affected by excessive pressure. One 
example of this is called a bedsore. Bedsores, also called decubitis ulcers, 
are caused by constant, long-term, unrelieved pressure on certain body parts 
that are bony, reducing blood flow to the area and leading to necrosis (tissue 
death). Bedsores are most common in elderly patients who have debilitating 
conditions that cause them to be immobile. Most hospitals and long-term 
care facilities have the practice of turning the patients every few hours to 
prevent the incidence of bedsores. If left untreated by removal of necrotized 
tissue, bedsores can be fatal if they become infected. 


The skin can also be affected by pressure associated with rapid growth. A 
stretch mark results when the dermis is stretched beyond its limits of 
elasticity, as the skin stretches to accommodate the excess pressure. Stretch 
marks usually accompany rapid weight gain during puberty and pregnancy. 
They initially have a reddish hue, but lighten over time. Other than for 
cosmetic reasons, treatment of stretch marks is not required. They occur 
most commonly over the hips and abdomen. 


Calluses 


When you wear shoes that do not fit well and are a constant source of 
abrasion on your toes, you tend to form a callus at the point of contact. This 
occurs because the basal stem cells in the stratum basale are triggered to 
divide more often to increase the thickness of the skin at the point of 
abrasion to protect the rest of the body from further damage. This is an 
example of a minor or local injury, and the skin manages to react and treat 
the problem independent of the rest of the body. Calluses can also form on 
your fingers if they are subject to constant mechanical stress, such as long 
periods of writing, playing string instruments, or video games. A corn is a 
specialized form of callus. Corns form from abrasions on the skin that result 
from an elliptical-type motion. 


Chapter Review 


Skin cancer is a result of damage to the DNA of skin cells, often due to 
excessive exposure to UV radiation. Basal cell carcinoma and squamous 
cell carcinoma are highly curable, and arise from cells in the stratum basale 
and stratum spinosum, respectively. Melanoma is the most dangerous form 
of skin cancer, affecting melanocytes, which can spread/metastasize to other 
organs. Burns are an injury to the skin that occur as a result of exposure to 
extreme heat, radiation, or chemicals. First-degree and second-degree burns 
usually heal quickly, but third-degree burns can be fatal because they 
penetrate the full thickness of the skin. Scars occur when there is repair of 
skin damage. Fibroblasts generate scar tissue in the form of collagen, which 
forms a basket-weave pattern that looks different from normal skin. 


Bedsores and stretch marks are the result of excessive pressure on the skin 
and underlying tissue. Bedsores are characterized by necrosis of tissue due 
to immobility, whereas stretch marks result from rapid growth. Eczema is 
an allergic reaction that manifests as a rash, and acne results from clogged 
sebaceous glands. Eczema and acne are usually long-term skin conditions 
that may be treated successfully in mild cases. Calluses and corns are the 
result of abrasive pressure on the skin. 


Review Questions 


Exercise: 


Problem:In general, skin cancers 


a. are easily treatable and not a major health concern 
b. occur due to poor hygiene 

c. can be reduced by limiting exposure to the sun 

d. affect only the epidermis 


Solution: 


C 


Exercise: 


Problem: Bedsores 


a. can be treated with topical moisturizers 

b. can result from deep massages 

c. are preventable by eliminating pressure points 
d. are caused by dry skin 


Solution: 


C 
Exercise: 


Problem: 


An individual has spent too much time sun bathing. Not only is his 
skin painful to touch, but small blisters have appeared in the affected 
area. This indicates that he has damaged which layers of his skin? 


a. epidermis only 

b. hypodermis only 

c. epidermis and hypodermis 
d. epidermis and dermis 


Solution: 


D 
Exercise: 
Problem: 
After a skin injury, the body initiates a wound-healing response. The 


first step of this response is the formation of a blood clot to stop 
bleeding. Which of the following would be the next response? 


a. increased production of melanin by melanocytes 

b. increased production of connective tissue 

c. an increase in Pacinian corpuscles around the wound 
d. an increased activity in the stratum lucidum 


Solution: 


B 
Exercise: 
Problem: 
Squamous cell carcinomas are the second most common of the skin 


cancers and are capable of metastasizing if not treated. This cancer 
affects which cells? 


a. basal cells of the stratum basale 

b. melanocytes of the stratum basale 

c. keratinocytes of the stratum spinosum 
d. Langerhans cells of the stratum lucidum 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem:Why do teenagers often experience acne? 


Solution: 


Acne results from a blockage of sebaceous glands by sebum. The 
blockage causes blackheads to form, which are susceptible to 
infection. The infected tissue then becomes red and inflamed. 
Teenagers experience this at high rates because the sebaceous glands 
become active during puberty. Hormones that are especially active 
during puberty stimulate the release of sebum, leading in many cases 
to blockages. 


Exercise: 


Problem:Why do scars look different from surrounding skin? 


Solution: 


Scars are made of collagen and do not have the cellular structure of 
normal skin. The tissue is fibrous and does not allow for the 
regeneration of accessory structures, such as hair follicles, and sweat 
or sebaceous glands. 


References 


American Cancer Society (US). Skin cancer: basal and squamous cell 
[Internet]. c2013 [cited 2012 Nov 1]. Available from: 
http://www.cancer.org/acs/groups/cid/documents/webcontent/003139- 
pdt.pdf. 


Lucile Packard Children’s Hospital at Stanford (US). Classification and 
treatment of burns [Internet]. Palo Alto (CA). c2012 [cited 2012 Nov 1]. 


Available from: 


Mayo Clinic (US). Basal cell carcinoma [Internet]. Scottsdale (AZ); c2012 
[cited 2012 Nov 1]. Available from: 
http://www.mayoclinic.com/health/basal-cell- 
carcinoma/ds00925/dsection=treatments-and-drugs. 


Beck, J. FYI: how much can a human body sweat before it runs out? 
Popular Science [Internet]. New York (NY); c2012 [cited 2012 Nov 1]. 


much-can-human-body-sweat-it-runs-out. 


Skin Cancer Foundation (US). Skin cancer facts [Internet]. New York (NY); 
c2013 [cited 2012 Nov 1]. Available from: http://www.skincancer.org/skin- 
cancer-information/skin-cancer-facts#top. 


Glossary 


acne 
skin condition due to infected sebaceous glands 


basal cell carcinoma 
cancer that originates from basal cells in the epidermis of the skin 


bedsore 
sore on the skin that develops when regions of the body start 
necrotizing due to constant pressure and lack of blood supply; also 
called decubitis ulcers 


callus 
thickened area of skin that arises due to constant abrasion 


corm 
type of callus that is named for its shape and the elliptical motion of 
the abrasive force 


eczema 


skin condition due to an allergic reaction, which resembles a rash 


first-degree burn 
superficial burn that injures only the epidermis 


fourth-degree burn 
burn in which full thickness of the skin and underlying muscle and 
bone is damaged 


keloid 
type of scar that has layers raised above the skin surface 


melanoma 
type of skin cancer that originates from the melanocytes of the skin 


metastasis 
spread of cancer cells from a source to other parts of the body 


scar 
collagen-rich skin formed after the process of wound healing that is 
different from normal skin 


second-degree burn 
partial-thickness burn that injures the epidermis and a portion of the 
dermis 


squamous cell carcinoma 
type of skin cancer that originates from the stratum spinosum of the 
epidermis 


stretch mark 
mark formed on the skin due to a sudden growth spurt and expansion 
of the dermis beyond its elastic limits 


third-degree burn 
burn that penetrates and destroys the full thickness of the skin 
(epidermis and dermis) 


The Functions of the Skeletal System 
By the end of this section, you will be able to: 


¢ Define bone, cartilage, and the skeletal system 
e List and describe the functions of the skeletal system 


Bone, or osseous tissue, is a hard, dense connective tissue that forms most 
of the adult skeleton, the support structure of the body. In the areas of the 
skeleton where bones move (for example, the ribcage and joints), cartilage, 
a semi-rigid form of connective tissue, provides flexibility and smooth 
surfaces for movement. The skeletal system is the body system composed 
of bones and cartilage and performs the following critical functions for the 
human body: 


e supports the body 

e facilitates movement 

¢ protects internal organs 

e produces blood cells 

e stores and releases minerals and fat 


Support, Movement, and Protection 


The most apparent functions of the skeletal system are the gross functions 
—those visible by observation. Simply by looking at a person, you can see 
how the bones support, facilitate movement, and protect the human body. 


Just as the steel beams of a building provide a scaffold to support its weight, 
the bones and cartilage of your skeletal system compose the scaffold that 
supports the rest of your body. Without the skeletal system, you would be a 
limp mass of organs, muscle, and skin. 


Bones also facilitate movement by serving as points of attachment for your 
muscles. While some bones only serve as a support for the muscles, others 
also transmit the forces produced when your muscles contract. From a 
mechanical point of view, bones act as levers and joints serve as fulcrums 
({link]). Unless a muscle spans a joint and contracts, a bone is not going to 


move. For information on the interaction of the skeletal and muscular 
systems, that is, the musculoskeletal system, seek additional content. 
Bones Support Movement 


Bones act as levers when 
muscles span a joint and 
contract. (credit: Benjamin J. 
DeLong) 


Bones also protect internal organs from injury by covering or surrounding 
them. For example, your ribs protect your lungs and heart, the bones of 
your vertebral column (spine) protect your spinal cord, and the bones of 
your cranium (skull) protect your brain ({link]). 

Bones Protect Brain 


The cranium completely 
surrounds and protects 
the brain from non- 
traumatic injury. 


Note: 

Career Connection 

Orthopedist 

An orthopedist is a doctor who specializes in diagnosing and treating 
disorders and injuries related to the musculoskeletal system. Some 
orthopedic problems can be treated with medications, exercises, braces, 
and other devices, but others may be best treated with surgery ([link]). 
Arm Brace 


An orthopedist will sometimes prescribe 
the use of a brace that reinforces the 
underlying bone structure it is being used 
to support. (credit: Juhan Sonin) 


While the origin of the word “orthopedics” (ortho- = “straight”; paed- = 
“child”), literally means “straightening of the child,” orthopedists can have 
patients who range from pediatric to geriatric. In recent years, orthopedists 
have even performed prenatal surgery to correct spina bifida, a congenital 
defect in which the neural canal in the spine of the fetus fails to close 
completely during embryologic development. 

Orthopedists commonly treat bone and joint injuries but they also treat 
other bone conditions including curvature of the spine. Lateral curvatures 
(scoliosis) can be severe enough to slip under the shoulder blade (scapula) 
forcing it up as a hump. Spinal curvatures can also be excessive 
dorsoventrally (kyphosis) causing a hunch back and thoracic compression. 
These curvatures often appear in preteens as the result of poor posture, 
abnormal growth, or indeterminate causes. Mostly, they are readily treated 
by orthopedists. As people age, accumulated spinal column injuries and 
diseases like osteoporosis can also lead to curvatures of the spine, hence 
the stooping you sometimes see in the elderly. 


Some orthopedists sub-specialize in sports medicine, which addresses both 
simple injuries, such as a sprained ankle, and complex injuries, such as a 
torn rotator cuff in the shoulder. Treatment can range from exercise to 
surgery. 


Mineral Storage, Energy Storage, and Hematopoiesis 


On a metabolic level, bone tissue performs several critical functions. For 
one, the bone matrix acts as a reservoir for a number of minerals important 
to the functioning of the body, especially calcium, and phosphorus. These 
minerals, incorporated into bone tissue, can be released back into the 
bloodstream to maintain levels needed to support physiological processes. 
Calcium ions, for example, are essential for muscle contractions and 
controlling the flow of other ions involved in the transmission of nerve 
impulses. 


Bone also serves as a site for fat storage and blood cell production. The 
softer connective tissue that fills the interior of most bone is referred to as 
bone marrow ((link]). There are two types of bone marrow: yellow marrow 
and red marrow. Yellow marrow contains adipose tissue; the triglycerides 
stored in the adipocytes of the tissue can serve as a source of energy. Red 
marrow is where hematopoiesis—the production of blood cells—takes 
place. Red blood cells, white blood cells, and platelets are all produced in 
the red marrow. 

Head of Femur Showing Red and Yellow Marrow 


Outer surface of bone 


Red marrow 


Yellow marrow 


The head of the femur contains both 
yellow and red marrow. Yellow marrow 
stores fat. Red marrow is responsible for 

hematopoiesis. (credit: modification of 
work by “stevenfruitsmaak”/Wikimedia 
Commons) 


Chapter Review 


The major functions of the bones are body support, facilitation of 
movement, protection of internal organs, storage of minerals and fat, and 
hematopoiesis. Together, the muscular system and skeletal system are 
known as the musculoskeletal system. 


Review Questions 


Exercise: 


Problem: 


Which function of the skeletal system would be especially important if 
you were in a car accident? 


a. storage of minerals 

b. protection of internal organs 
c. facilitation of movement 

d. fat storage 


Solution: 


B 


Exercise: 


Problem:Bone tissue can be described as 


a. dead calcified tissue 

b. cartilage 

c. the skeletal system 

d. dense, hard connective tissue 


Solution: 


D 


Exercise: 


Problem: Without red marrow, bones would not be able to 


a. store phosphate 
b. store calcium 

c. make blood cells 
d. move like levers 


Solution: 
C 
Exercise: 
Problem: Yellow marrow has been identified as 
a. an area of fat storage 
b. a point of attachment for muscles 


c. the hard portion of bone 
d. the cause of kyphosis 


Solution: 


A 
Exercise: 
Problem: Which of the following can be found in areas of movement? 


a. hematopoiesis 
b. cartilage 

c. yellow marrow 
d. red marrow 


Solution: 
B 
Exercise: 


Problem:The skeletal system is made of 


a. muscles and tendons 
b. bones and cartilage 
c. vitreous humor 

d. minerals and fat 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


The skeletal system is composed of bone and cartilage and has many 
functions. Choose three of these functions and discuss what features of 
the skeletal system allow it to accomplish these functions. 


Solution: 


It supports the body. The rigid, yet flexible skeleton acts as a 
framework to support the other organs of the body. 


It facilitates movement. The movable joints allow the skeleton to 
change shape and positions; that is, move. 


It protects internal organs. Parts of the skeleton enclose or partly 
enclose various organs of the body including our brain, ears, heart, and 
lungs. Any trauma to these organs has to be mediated through the 
skeletal system. 


It produces blood cells. The central cavity of long bones is filled with 
marrow. The red marrow is responsible for forming red and white 
blood cells. 


It stores and releases minerals and fat. The mineral component of 
bone, in addition to providing hardness to bone, provides a mineral 
reservoir that can be tapped as needed. Additionally, the yellow 
marrow, which is found in the central cavity of long bones along with 
red marrow, serves as a Storage site for fat. 


Glossary 


bone 
hard, dense connective tissue that forms the structural elements of the 
skeleton 


cartilage 


semi-rigid connective tissue found on the skeleton in areas where 
flexibility and smooth surfaces support movement 


hematopoiesis 
production of blood cells, which occurs in the red marrow of the bones 


orthopedist 
doctor who specializes in diagnosing and treating musculoskeletal 
disorders and injuries 


osseous tissue 
bone tissue; a hard, dense connective tissue that forms the structural 
elements of the skeleton 


red marrow 
connective tissue in the interior cavity of a bone where hematopoiesis 
takes place 


skeletal system 
organ system composed of bones and cartilage that provides for 
movement, support, and protection 


yellow marrow 
connective tissue in the interior cavity of a bone where fat is stored 


Bone Classification 
By the end of this section, you will be able to: 


¢ Classify bones according to their shapes 
e Describe the function of each category of bones 


The 206 bones that compose the adult skeleton are divided into five 
categories based on their shapes ((link]). Their shapes and their functions 
are related such that each categorical shape of bone has a distinct function. 
Classifications of Bones 


Irregular bone 


Flat bone 


Vertebra 


Sternum 


Long bone 


Short bones 


Lateral 
cuneiform 


Sesamoid bone 


Intermediate 
cuneiform 


= - Medial 
- y cuneiform 


Patella ne - 


Bones are classified according to their shape. 


Long Bones 


A long bone is one that is cylindrical in shape, being longer than it is wide. 
Keep in mind, however, that the term describes the shape of a bone, not its 
size. Long bones are found in the arms (humerus, ulna, radius) and legs 
(femur, tibia, fibula), as well as in the fingers (metacarpals, phalanges) and 
toes (metatarsals, phalanges). Long bones function as levers; they move 
when muscles contract. 


Short Bones 


A short bone is one that is cube-like in shape, being approximately equal in 
length, width, and thickness. The only short bones in the human skeleton 
are in the carpals of the wrists and the tarsals of the ankles. Short bones 
provide stability and support as well as some limited motion. 


Flat Bones 


The term “flat bone” is somewhat of a misnomer because, although a flat 
bone is typically thin, it is also often curved. Examples include the cranial 
(skull) bones, the scapulae (shoulder blades), the sternum (breastbone), and 
the ribs. Flat bones serve as points of attachment for muscles and often 
protect internal organs. 


Irregular Bones 


An irregular bone is one that does not have any easily characterized shape 
and therefore does not fit any other classification. These bones tend to have 
more complex shapes, like the vertebrae that support the spinal cord and 
protect it from compressive forces. Many facial bones, particularly the ones 
containing sinuses, are classified as irregular bones. 


Sesamoid Bones 


A sesamoid bone is a small, round bone that, as the name suggests, is 
shaped like a sesame seed. These bones form in tendons (the sheaths of 
tissue that connect bones to muscles) where a great deal of pressure is 
generated in a joint. The sesamoid bones protect tendons by helping them 


overcome compressive forces. Sesamoid bones vary in number and 
placement from person to person but are typically found in tendons 
associated with the feet, hands, and knees. The patellae (singular = patella) 
are the only sesamoid bones found in common with every person. [link] 
reviews bone classifications with their associated features, functions, and 


examples. 


Bone Classifications 


Bone 


classification 


Long 


Short 


Features 


Cylinder-like 
shape, longer 
than it is wide 


Cube-like 
shape, 
approximately 
equal in 
length, width, 
and thickness 


Function(s) 


Leverage 


Provide 
stability, 
support, 
while 
allowing for 
some 
motion 


Examples 


Femur, tibia, 
fibula, 
metatarsals, 
humerus, 
ulna, radius, 
metacarpals, 
phalanges 


Carpals, 
tarsals 


Bone Classifications 


Bone 
classification Features Function(s) Examples 
Points of 
attachment Sternum, 
Flat Thin and for muscles; ribs, 
curved protectors scapulae, 
of internal cranial bones 
organs 
Pr 
Complex — Vertebrae, 
Irregular internal 
shape facial bones 
organs 
Pr 
Small and oe 
fount: tendons 
i : : from Patell 
Sesamoid anbeddedan 0) atellae 
compressive 
tendons 
forces 
Chapter Review 


Bones can be classified according to their shapes. Long bones, such as the 
femur, are longer than they are wide. Short bones, such as the carpals, are 
approximately equal in length, width, and thickness. Flat bones are thin, but 
are often curved, such as the ribs. Irregular bones such as those of the face 
have no characteristic shape. Sesamoid bones, such as the patellae, are 
small and round, and are located in tendons. 


Review Questions 


Exercise: 


Problem: 


Most of the bones of the arms and hands are long bones; however, the 
bones in the wrist are categorized as 


a. flat bones 

b. short bones 

c. sesamoid bones 
d. irregular bones 


Solution: 


B 


Exercise: 


Problem:Sesamoid bones are found embedded in 


a. joints 

b. muscles 
c. ligaments 
d. tendons 


Solution: 


D 
Exercise: 


Problem: 
Bones that surround the spinal cord are classified as bones. 


a. irregular 
b. sesamoid 
c. flat 

d. short 


Solution: 


A 
Exercise: 


Problem: 


Which category of bone is among the most numerous in the skeleton? 
a. long bone 
b. sesamoid bone 


c. short bone 
d. flat bone 


Solution: 


A 


Exercise: 


Problem:Long bones enable body movement by acting as a 
a. counterweight 
b. resistive force 
c. lever 
d. fulcrum 


Solution: 


GC 


Critical Thinking Questions 


Exercise: 


Problem: 


What are the structural and functional differences between a tarsal and 
a metatarsal? 


Solution: 


Structurally, a tarsal is a short bone, meaning its length, width, and 
thickness are about equal, while a metatarsal is a long bone whose 
length is greater than its width. Functionally, the tarsal provides 
limited motion, while the metatarsal acts as a lever. 


Exercise: 
Problem: 


What are the structural and functional differences between the femur 
and the patella? 


Solution: 


Structurally, the femur is a long bone, meaning its length is greater 
than its width, while the patella, a sesamoid bone, is small and round. 
Functionally, the femur acts as a lever, while the patella protects the 
patellar tendon from compressive forces. 


Glossary 


flat bone 
thin and curved bone; serves as a point of attachment for muscles and 
protects internal organs 


irregular bone 
bone of complex shape; protects internal organs from compressive 
forces 


long bone 
cylinder-shaped bone that is longer than it is wide; functions as a lever 


sesamoid bone 
small, round bone embedded in a tendon; protects the tendon from 
compressive forces 


short bone 
cube-shaped bone that is approximately equal in length, width, and 
thickness; provides limited motion 


Bone Structure 
By the end of this section, you will be able to: 


e Identify the anatomical features of a bone 

¢ Define and list examples of bone markings 

¢ Describe the histology of bone tissue 

e Compare and contrast compact and spongy bone 

e Identify the structures that compose compact and spongy bone 
e Describe how bones are nourished and innervated 


Bone tissue (osseous tissue) differs greatly from other tissues in the body. 
Bone is hard and many of its functions depend on that characteristic 
hardness. Later discussions in this chapter will show that bone is also 
dynamic in that its shape adjusts to accommodate stresses. This section will 
examine the gross anatomy of bone first and then move on to its histology. 


Gross Anatomy of Bone 


The structure of a long bone allows for the best visualization of all of the 
parts of a bone ([link]). A long bone has two parts: the diaphysis and the 
epiphysis. The diaphysis is the tubular shaft that runs between the proximal 
and distal ends of the bone. The hollow region in the diaphysis is called the 
medullary cavity, which is filled with yellow marrow. The walls of the 
diaphysis are composed of dense and hard compact bone. 

Anatomy of a Long Bone 


Articular cartilage 


Proximal 
epiphysis 


Metaphysis Spongy bone 
Epiphyseal line 
Red bone marrow 


Endosteum 


Compact bone 


Medullary cavity 
Diaphysis Yellow bone marrow 


Periosteum 


Nutrient artery 


Distal 
epiphysis 


Articular cartilage 


A typical long bone shows the 
gross anatomical characteristics of 
bone. 


The wider section at each end of the bone is called the epiphysis (plural = 
epiphyses), which is filled with spongy bone. Red marrow fills the spaces in 
the spongy bone. Each epiphysis meets the diaphysis at the metaphysis, the 
narrow area that contains the epiphyseal plate (growth plate), a layer of 
hyaline (transparent) cartilage in a growing bone. When the bone stops 
growing in early adulthood (approximately 18-21 years), the cartilage is 


replaced by osseous tissue and the epiphyseal plate becomes an epiphyseal 
line. 


The medullary cavity has a delicate membranous lining called the 
endosteum (end- = “inside”; oste- = “bone”), where bone growth, repair, 
and remodeling occur. The outer surface of the bone is covered with a 
fibrous membrane called the periosteum (peri- = “around” or 
“surrounding”). The periosteum contains blood vessels, nerves, and 
lymphatic vessels that nourish compact bone. Tendons and ligaments also 
attach to bones at the periosteum. The periosteum covers the entire outer 
surface except where the epiphyses meet other bones to form joints ([link]). 
In this region, the epiphyses are covered with articular cartilage, a thin 
layer of cartilage that reduces friction and acts as a shock absorber. 
Periosteum and Endosteum 


Periosteum / 


Endosteum 


Periosteum Osteoclast 


(fibrous layer) Osteocyte 


in lacuna Bone matrix 
Osteocyte 


Periosteum Osteogenic cell 


(cellular layer) Osteoblast 


The periosteum forms the outer surface of bone, and 
the endosteum lines the medullary cavity. 


Flat bones, like those of the cranium, consist of a layer of diploé (spongy 
bone), lined on either side by a layer of compact bone ([link]). The two 
layers of compact bone and the interior spongy bone work together to 
protect the internal organs. If the outer layer of a cranial bone fractures, the 
brain is still protected by the intact inner layer. 

Anatomy of a Flat Bone 


| \i 
4 


‘i ten as 
Periosteum RN Ce I ee 
~ ° Dee | c=! 


OSC” bone (diploé) 


This cross-section of a flat bone shows the 
spongy bone (diploé) lined on either side by a 
layer of compact bone. 


Bone Markings 


The surface features of bones vary considerably, depending on the function 
and location in the body. [link] describes the bone markings, which are 
illustrated in ([link]). There are three general classes of bone markings: (1) 
articulations, (2) projections, and (3) holes. As the name implies, an 
articulation is where two bone surfaces come together (articulus = “joint”). 
These surfaces tend to conform to one another, such as one being rounded 
and the other cupped, to facilitate the function of the articulation. A 
projection is an area of a bone that projects above the surface of the bone. 
These are the attachment points for tendons and ligaments. In general, their 
size and shape is an indication of the forces exerted through the attachment 
to the bone. A hole is an opening or groove in the bone that allows blood 
vessels and nerves to enter the bone. As with the other markings, their size 
and shape reflect the size of the vessels and nerves that penetrate the bone 
at these points. 


Bone Markings 


Marking 


Articulations 


Head 


Facet 


Condyle 


Projections 


Protuberance 


Process 


Spine 


Tubercle 


Tuberosity 


Description 


Where two 
bones meet 


Prominent 
rounded 
surface 


Flat surface 


Rounded 
surface 


Raised 
markings 


Protruding 


Prominence 
feature 


Sharp 
process 


Small, 
rounded 
process 


Rough 
surface 


Example 


Knee joint 


Head of femur 


Vertebrae 


Occipital condyles 


Spinous process of the vertebrae 


Chin 


Transverse process of vertebra 


Ischial spine 


Tubercle of humerus 


Deltoid tuberosity 


Bone Markings 


Marking 


Line 


Crest 


Holes 


Fossa 


Fovea 


Sulcus 


Canal 


Fissure 


Foramen 


Meatus 


Description 
Slight, 
elongated 
ridge 

Ridge 


Holes and 
depressions 


Elongated 
basin 


Small pit 


Groove 


Passage in 
bone 


Slit through 
bone 


Hole 
through 
bone 


Opening 
into canal 


Example 


Temporal lines of the parietal 
bones 


Iliac crest 


Foramen (holes through which 
blood vessels can pass through) 


Mandibular fossa 


Fovea capitis on the head of the 
femur 


Sigmoid sulcus of the temporal 
bones 


Auditory canal 


Auricular fissure 


Foramen magnum in the 
occipital bone 


External auditory meatus 


Bone Markings 


Marking Description Example 
Air-filled 

Sinus space in Nasal sinus 
bone 


Bone Features 


Examples of processes formed where Examples of an elevation or depression 
tendons or ligaments attach 


Fovea capitis 


17 Sulcus 


Head 
Tubercle Pay 
\ 


Tuberosity 


Pelvis 


Fossa 


Humerus Examples of openings 


Tubercle Condyle 
Facet 


Foramen 


Condyles 


Examples of processes formed to Canal , \ \ Fissure 
articulate with adjacent bones \ \ 


Protuberance 


Skull 


The surface features of bones depend on their function, 
location, attachment of ligaments and tendons, or the 
penetration of blood vessels and nerves. 


Bone Cells and Tissue 


Bone contains a relatively small number of cells entrenched in a matrix of 
collagen fibers that provide a surface for inorganic salt crystals to adhere. 
These salt crystals form when calcium phosphate and calcium carbonate 
combine to create hydroxyapatite, which incorporates other inorganic salts 
like magnesium hydroxide, fluoride, and sulfate as it crystallizes, or 
calcifies, on the collagen fibers. The hydroxyapatite crystals give bones 
their hardness and strength, while the collagen fibers give them flexibility 
so that they are not brittle. 


Although bone cells compose a small amount of the bone volume, they are 
crucial to the function of bones. Four types of cells are found within bone 
tissue: osteoblasts, osteocytes, osteogenic cells, and osteoclasts ([link]). 
Bone Cells 


wet &. 


Osteocyte Osteoblast Osteogenic cell Osteoclast 
(maintains (forms bone matrix) (stem cell) (resorbs bone) 
bone tissue) 


Four types of cells are found within bone 
tissue. Osteogenic cells are undifferentiated 
and develop into osteoblasts. When 
osteoblasts get trapped within the calcified 
matrix, their structure and function changes, 
and they become osteocytes. Osteoclasts 


develop from monocytes and macrophages 
and differ in appearance from other bone 
cells. 


The osteoblast is the bone cell responsible for forming new bone and is 
found in the growing portions of bone, including the periosteum and 
endosteum. Osteoblasts, which do not divide, synthesize and secrete the 
collagen matrix and calcium salts. As the secreted matrix surrounding the 
osteoblast calcifies, the osteoblast become trapped within it; as a result, it 
changes in structure and becomes an osteocyte, the primary cell of mature 
bone and the most common type of bone cell. Each osteocyte is located in a 
space called a lacuna and is surrounded by bone tissue. Osteocytes 
maintain the mineral concentration of the matrix via the secretion of 
enzymes. Like osteoblasts, osteocytes lack mitotic activity. They can 
communicate with each other and receive nutrients via long cytoplasmic 
processes that extend through canaliculi (singular = canaliculus), channels 
within the bone matrix. 


If osteoblasts and osteocytes are incapable of mitosis, then how are they 
replenished when old ones die? The answer lies in the properties of a third 
category of bone cells—the osteogenic cell. These osteogenic cells are 
undifferentiated with high mitotic activity and they are the only bone cells 
that divide. Immature osteogenic cells are found in the deep layers of the 
periosteum and the marrow. They differentiate and develop into osteoblasts. 


The dynamic nature of bone means that new tissue is constantly formed, 
and old, injured, or unnecessary bone is dissolved for repair or for calcium 
release. The cell responsible for bone resorption, or breakdown, is the 
osteoclast. They are found on bone surfaces, are multinucleated, and 
originate from monocytes and macrophages, two types of white blood cells, 
not from osteogenic cells. Osteoclasts are continually breaking down old 
bone while osteoblasts are continually forming new bone. The ongoing 
balance between osteoblasts and osteoclasts is responsible for the constant 
but subtle reshaping of bone. [link] reviews the bone cells, their functions, 
and locations. 


Bone Cells 


Cell type Function 
Osteogenic Develop into 
cells osteoblasts 
Osteoblasts Bone formation 
Maintain mineral 
Osteocytes concentration of 
matrix 
Osteoclasts Bone resorption 
Compact and Spongy Bone 


Location 


Deep layers of the 
periosteum and the marrow 


Growing portions of bone, 
including periosteum and 
endosteum 


Entrapped in matrix 


Bone surfaces and at sites of 
old, injured, or unneeded 
bone 


The differences between compact and spongy bone are best explored via 
their histology. Most bones contain compact and spongy osseous tissue, but 
their distribution and concentration vary based on the bone’s overall 
function. Compact bone is dense so that it can withstand compressive 
forces, while spongy (cancellous) bone has open spaces and supports shifts 
in weight distribution. 


Compact Bone 


Compact bone is the denser, stronger of the two types of bone tissue 
({link]). It can be found under the periosteum and in the diaphyses of long 
bones, where it provides support and protection. 

Diagram of Compact Bone 


(a) This cross-sectional view of compact bone shows the basic 
structural unit, the osteon. (b) In this micrograph of the osteon, you can 
clearly see the concentric lamellae and central canals. LM x 40. 
(Micrograph provided by the Regents of University of Michigan 
Medical School © 2012) 


Compact bone Spongy bone 


Medullary cavity 


Periosteum 


Concentric lamellae 


Osteon 


Lymphatic vessel 
Circumferential 


lamellae Nerve 


Periosteal artery Blood vessels 
Periosteal vein rabeéulae 


Periosteum: 
Outer fibrous layer 
Inner osteogenic layer 


Interstitial lamellae Medullar cavity 


Perforating canal 
Spongy bone 


Central canal 
Blood vessels 
Lymphatic vessel 


Nerve 
Compact bone 


The microscopic structural unit of compact bone is called an osteon, or 
Haversian system. Each osteon is composed of concentric rings of calcified 
matrix called lamellae (singular = lamella). Running down the center of 
each osteon is the central canal, or Haversian canal, which contains blood 
vessels, nerves, and lymphatic vessels. These vessels and nerves branch off 


at right angles through a perforating canal, also known as Volkmann’s 
canals, to extend to the periosteum and endosteum. 


The osteocytes are located inside spaces called lacunae (singular = lacuna), 
found at the borders of adjacent lamellae. As described earlier, canaliculi 
connect with the canaliculi of other lacunae and eventually with the central 
canal. This system allows nutrients to be transported to the osteocytes and 
wastes to be removed from them. 


Spongy (Cancellous) Bone 


Like compact bone, spongy bone, also known as cancellous bone, contains 
osteocytes housed in lacunae, but they are not arranged in concentric 
circles. Instead, the lacunae and osteocytes are found in a lattice-like 
network of matrix spikes called trabeculae (singular = trabecula) ([link]). 
The trabeculae may appear to be a random network, but each trabecula 
forms along lines of stress to provide strength to the bone. The spaces of the 
trabeculated network provide balance to the dense and heavy compact bone 
by making bones lighter so that muscles can move them more easily. In 
addition, the spaces in some spongy bones contain red marrow, protected by 
the trabeculae, where hematopoiesis occurs. 


Osteoclast 


Osteoblasts aligned 
along trabeculae of 
new bone 


Canaliculi Endosteum _Lamellae 
openings : Ge Canaliculi 
on surface aay 

Lamellae 


Spongy bone is composed of trabeculae that contain 
the osteocytes. Red marrow fills the spaces in some 
bones. 


Note: 

Aging and the... 

Skeletal System: Paget’s Disease 

Paget’s disease usually occurs in adults over age 40. It is a disorder of the 
bone remodeling process that begins with overactive osteoclasts. This 
means more bone is resorbed than is laid down. The osteoblasts try to 
compensate but the new bone they lay down is weak and brittle and 
therefore prone to fracture. 

While some people with Paget’s disease have no symptoms, others 
experience pain, bone fractures, and bone deformities ([link]). Bones of the 
pelvis, skull, spine, and legs are the most commonly affected. When 
occurring in the skull, Paget’s disease can cause headaches and hearing 
loss. 


Paget's Disease 


Normal Paget’s disease 


Normal leg bones are relatively 
Straight, but those affected by 
Paget’s disease are porous and 

curved. 


What causes the osteoclasts to become overactive? The answer is still 
unknown, but hereditary factors seem to play a role. Some scientists 
believe Paget’s disease is due to an as-yet-unidentified virus. 

Paget’s disease is diagnosed via imaging studies and lab tests. X-rays may 
show bone deformities or areas of bone resorption. Bone scans are also 
useful. In these studies, a dye containing a radioactive ion is injected into 
the body. Areas of bone resorption have an affinity for the ion, so they will 
light up on the scan if the ions are absorbed. In addition, blood levels of an 
enzyme called alkaline phosphatase are typically elevated in people with 
Paget’s disease. 

Bisphosphonates, drugs that decrease the activity of osteoclasts, are often 
used in the treatment of Paget’s disease. However, in a small percentage of 
cases, bisphosphonates themselves have been linked to an increased risk of 
fractures because the old bone that is left after bisphosphonates are 
administered becomes worn out and brittle. Still, most doctors feel that the 
benefits of bisphosphonates more than outweigh the risk; the medical 
professional has to weigh the benefits and risks on a case-by-case basis. 
Bisphosphonate treatment can reduce the overall risk of deformities or 
fractures, which in turn reduces the risk of surgical repair and its associated 
risks and complications. 


Blood and Nerve Supply 


The spongy bone and medullary cavity receive nourishment from arteries 
that pass through the compact bone. The arteries enter through the nutrient 
foramen (plural = foramina), small openings in the diaphysis ((link]). The 
osteocytes in spongy bone are nourished by blood vessels of the periosteum 
that penetrate spongy bone and blood that circulates in the marrow cavities. 


As the blood passes through the marrow cavities, it is collected by veins, 
which then pass out of the bone through the foramina. 


In addition to the blood vessels, nerves follow the same paths into the bone 
where they tend to concentrate in the more metabolically active regions of 
the bone. The nerves sense pain, and it appears the nerves also play roles in 
regulating blood supplies and in bone growth, hence their concentrations in 
metabolically active sites of the bone. 

Diagram of Blood and Nerve Supply to Bone 


ere Articular 
—_ ae zm) cartilage 


Epiphyseal 
artery 
and vein 


= Metaphyseal 


artery 
and vein 


Periosteum 


Compact 
bone 


Nutrient 
artery 
and vein 


Nutrient 


foramen Medullary cavity 


Metaphyseal 
artery and vein 


Metaphysis —|| 


Epiphyseal 
line 


Blood vessels and nerves 
enter the bone through the 
nutrient foramen. 


Note: 


Watch this video to see the microscopic features of a bone. 


Chapter Review 


A hollow medullary cavity filled with yellow marrow runs the length of the 
diaphysis of a long bone. The walls of the diaphysis are compact bone. The 
epiphyses, which are wider sections at each end of a long bone, are filled 
with spongy bone and red marrow. The epiphyseal plate, a layer of hyaline 
cartilage, is replaced by osseous tissue as the organ grows in length. The 
medullary cavity has a delicate membranous lining called the endosteum. 
The outer surface of bone, except in regions covered with articular 
cartilage, is covered with a fibrous membrane called the periosteum. Flat 
bones consist of two layers of compact bone surrounding a layer of spongy 
bone. Bone markings depend on the function and location of bones. 
Articulations are places where two bones meet. Projections stick out from 
the surface of the bone and provide attachment points for tendons and 
ligaments. Holes are openings or depressions in the bones. 


Bone matrix consists of collagen fibers and organic ground substance, 
primarily hydroxyapatite formed from calcium salts. Osteogenic cells 
develop into osteoblasts. Osteoblasts are cells that make new bone. They 
become osteocytes, the cells of mature bone, when they get trapped in the 
matrix. Osteoclasts engage in bone resorption. Compact bone is dense and 
composed of osteons, while spongy bone is less dense and made up of 
trabeculae. Blood vessels and nerves enter the bone through the nutrient 
foramina to nourish and innervate bones. 


Review Questions 


Exercise: 


Problem: 


Which of the following occurs in the spongy bone of the epiphysis? 


a. bone growth 

b. bone remodeling 
c. hematopoiesis 

d. shock absorption 


Solution: 


C 


Exercise: 


Problem:The diaphysis contains 


a. the metaphysis 
b. fat stores 

c. spongy bone 
d. compact bone 


Solution: 


B 
Exercise: 


Problem: 


The fibrous membrane covering the outer surface of the bone is the 


a. periosteum 
b. epiphysis 
c. endosteum 


d. diaphysis 


Solution: 


A 


Exercise: 


Problem: Which of the following are incapable of undergoing mitosis? 


a. osteoblasts and osteoclasts 
b. osteocytes and osteoclasts 
c. osteoblasts and osteocytes 
d. osteogenic cells and osteoclasts 


Solution: 


‘§ 


Exercise: 


Problem: Which cells do not originate from osteogenic cells? 


a. osteoblasts 
b. osteoclasts 
c. osteocytes 
d. osteoprogenitor cells 


Solution: 


D 


Exercise: 


Problem: 


Which of the following are found in compact bone and cancellous 
bone? 


a. Haversian systems 
b. Haversian canals 
c. lamellae 

d. lacunae 


Solution: 


C 


Exercise: 


Problem: Which of the following are only found in cancellous bone? 


a. canaliculi 

b. Volkmann’s canals 
c. trabeculae 

d. calcium salts 


Solution: 


C 
Exercise: 
Problem: 


The area of a bone where the nutrient foramen passes forms what kind 
of bone marking? 


a. a hole 
b. a facet 
c. a canal 


d. a fissure 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: 


If the articular cartilage at the end of one of your long bones were to 
degenerate, what symptoms do you think you would experience? 
Why? 


Solution: 


If the articular cartilage at the end of one of your long bones were to 
deteriorate, which is actually what happens in osteoarthritis, you would 
experience joint pain at the end of that bone and limitation of motion at 
that joint because there would be no cartilage to reduce friction 
between adjacent bones and there would be no cartilage to act as a 
shock absorber. 


Exercise: 
Problem: 


In what ways is the structural makeup of compact and spongy bone 
well suited to their respective functions? 


Solution: 


The densely packed concentric rings of matrix in compact bone are 
ideal for resisting compressive forces, which is the function of 
compact bone. The open spaces of the trabeculated network of spongy 


bone allow spongy bone to support shifts in weight distribution, which 
is the function of spongy bone. 


Glossary 


articular cartilage 
thin layer of cartilage covering an epiphysis; reduces friction and acts 
as a shock absorber 


articulation 
where two bone surfaces meet 


canaliculi 
(singular = canaliculus) channels within the bone matrix that house 
one of an osteocyte’s many cytoplasmic extensions that it uses to 
communicate and receive nutrients 


central canal 
longitudinal channel in the center of each osteon; contains blood 
vessels, nerves, and lymphatic vessels; also known as the Haversian 
canal 


compact bone 
dense osseous tissue that can withstand compressive forces 


diaphysis 
tubular shaft that runs between the proximal and distal ends of a long 
bone 


diploé 
layer of spongy bone, that is sandwiched between two the layers of 


compact bone found in flat bones 


endosteum 
delicate membranous lining of a bone’s medullary cavity 


epiphyseal plate 


(also, growth plate) sheet of hyaline cartilage in the metaphysis of an 
immature bone; replaced by bone tissue as the organ grows in length 


epiphysis 
wide section at each end of a long bone; filled with spongy bone and 
red marrow 


hole 
opening or depression in a bone 


lacunae 
(singular = lacuna) spaces in a bone that house an osteocyte 


medullary cavity 
hollow region of the diaphysis; filled with yellow marrow 


nutrient foramen 
small opening in the middle of the external surface of the diaphysis, 
through which an artery enters the bone to provide nourishment 


osteoblast 
cell responsible for forming new bone 


osteoclast 
cell responsible for resorbing bone 


osteocyte 
primary cell in mature bone; responsible for maintaining the matrix 


osteogenic cell 
undifferentiated cell with high mitotic activity; the only bone cells that 
divide; they differentiate and develop into osteoblasts 


osteon 
(also, Haversian system) basic structural unit of compact bone; made 
of concentric layers of calcified matrix 


perforating canal 


(also, Volkmann’s canal) channel that branches off from the central 
canal and houses vessels and nerves that extend to the periosteum and 
endosteum 


periosteum 
fibrous membrane covering the outer surface of bone and continuous 
with ligaments 


projection 
bone markings where part of the surface sticks out above the rest of the 
surface, where tendons and ligaments attach 


spongy bone 
(also, cancellous bone) trabeculated osseous tissue that supports shifts 
in weight distribution 


trabeculae 
(singular = trabecula) spikes or sections of the lattice-like matrix in 
spongy bone 


Bone Formation and Development 
By the end of this section, you will be able to: 


e Explain the function of cartilage 

e List the steps of intramembranous ossification 

e List the steps of endochondral ossification 

e Explain the growth activity at the epiphyseal plate 

e Compare and contrast the processes of modeling and remodeling 


In the early stages of embryonic development, the embryo’s skeleton 
consists of fibrous membranes and hyaline cartilage. By the sixth or seventh 
week of embryonic life, the actual process of bone development, 
ossification (osteogenesis), begins. There are two osteogenic pathways— 
intramembranous ossification and endochondral ossification—but bone is 
the same regardless of the pathway that produces it. 


Cartilage Templates 


Bone is a replacement tissue; that is, it uses a model tissue on which to lay 
down its mineral matrix. For skeletal development, the most common 
template is cartilage. During fetal development, a framework is laid down 
that determines where bones will form. This framework is a flexible, semi- 
solid matrix produced by chondroblasts and consists of hyaluronic acid, 
chondroitin sulfate, collagen fibers, and water. As the matrix surrounds and 
isolates chondroblasts, they are called chondrocytes. Unlike most 
connective tissues, cartilage is avascular, meaning that it has no blood 
vessels supplying nutrients and removing metabolic wastes. All of these 
functions are carried on by diffusion through the matrix. This is why 
damaged cartilage does not repair itself as readily as most tissues do. 


Throughout fetal development and into childhood growth and development, 
bone forms on the cartilaginous matrix. By the time a fetus is born, most of 
the cartilage has been replaced with bone. Some additional cartilage will be 
replaced throughout childhood, and some cartilage remains in the adult 
skeleton. 


Intramembranous Ossification 


During intramembranous ossification, compact and spongy bone 
develops directly from sheets of mesenchymal (undifferentiated) connective 
tissue. The flat bones of the face, most of the cranial bones, and the 
clavicles (collarbones) are formed via intramembranous ossification. 


The process begins when mesenchymal cells in the embryonic skeleton 
gather together and begin to differentiate into specialized cells ({link]a). 
Some of these cells will differentiate into capillaries, while others will 
become osteogenic cells and then osteoblasts. Although they will ultimately 
be spread out by the formation of bone tissue, early osteoblasts appear in a 
cluster called an ossification center. 


The osteoblasts secrete osteoid, uncalcified matrix, which calcifies 
(hardens) within a few days as mineral salts are deposited on it, thereby 
entrapping the osteoblasts within. Once entrapped, the osteoblasts become 
osteocytes ([{link]b). As osteoblasts transform into osteocytes, osteogenic 
cells in the surrounding connective tissue differentiate into new osteoblasts. 


Osteoid (unmineralized bone matrix) secreted around the capillaries results 
in a trabecular matrix, while osteoblasts on the surface of the spongy bone 
become the periosteum ([link]c). The periosteum then creates a protective 
layer of compact bone superficial to the trabecular bone. The trabecular 
bone crowds nearby blood vessels, which eventually condense into red 
marrow ((link]d). 

Intramembranous Ossification 


Mesenchymal 
cells 
Osteoid 


Osteoblast 


Ossification 
center 


Osteocyte 


New bone 
matrix 
Osteoid 


Osteoblast 


Mesenchyme —= SS 


forms the ES ——— 
6. y 


-——_ Fibrous 


periosteum 
periosteum 


Osteoblast 


Compact bone 


Trabeculae 


Spongy bone 
(cavities contain 


Zz = — = = red marrow) 


Blood vessel 


Intramembranous ossification follows four steps. (a) 
Mesenchymal cells group into clusters, and ossification centers 
form. (b) Secreted osteoid traps osteoblasts, which then 
become osteocytes. (c) Trabecular matrix and periosteum form. 
(d) Compact bone develops superficial to the trabecular bone, 
and crowded blood vessels condense into red marrow. 


Intramembranous ossification begins in utero during fetal development and 
continues on into adolescence. At birth, the skull and clavicles are not fully 
ossified nor are the sutures of the skull closed. This allows the skull and 
shoulders to deform during passage through the birth canal. The last bones 
to ossify via intramembranous ossification are the flat bones of the face, 
which reach their adult size at the end of the adolescent growth spurt. 


Endochondral Ossification 


In endochondral ossification, bone develops by replacing hyaline 
cartilage. Cartilage does not become bone. Instead, cartilage serves as a 
template to be completely replaced by new bone. Endochondral ossification 
takes much longer than intramembranous ossification. Bones at the base of 
the skull and long bones form via endochondral ossification. 


In a long bone, for example, at about 6 to 8 weeks after conception, some of 
the mesenchymal cells differentiate into chondrocytes (cartilage cells) that 
form the cartilaginous skeletal precursor of the bones ([link]a). Soon after, 
the perichondrium, a membrane that covers the cartilage, appears [link]b). 
Endochondral Ossification 


Perichondrium 


Primary 
ossification 
center 


Hyaline 
cartilage 


Calcified 
matrix 


(c) Periosteum 
(covers compact 
bone) 


Medullary 
cavity 


Artery and vein 
(provide nutrients 
to bone) 


(d) 


Secondary 
ossification 
center 


Articular cartilage 
_—— Artery 
and vein 


Artery and vein 
(provide nutrients 
to bone) 


(e) 


Endochondral ossification follows five steps. (a) Mesenchymal cells 


differentiate into chondrocytes. (b) The cartilage model of the future 
bony skeleton and the perichondrium form. (c) Capillaries penetrate 
cartilage. Perichondrium transforms into periosteum. Periosteal collar 
develops. Primary ossification center develops. (d) Cartilage and 
chondrocytes continue to grow at ends of the bone. (e) Secondary 
ossification centers develop. (f) Cartilage remains at epiphyseal 
(growth) plate and at joint surface as articular cartilage. 


As more matrix is produced, the chondrocytes in the center of the 
cartilaginous model grow in size. As the matrix calcifies, nutrients can no 
longer reach the chondrocytes. This results in their death and the 
disintegration of the surrounding cartilage. Blood vessels invade the 
resulting spaces, not only enlarging the cavities but also carrying osteogenic 
cells with them, many of which will become osteoblasts. These enlarging 
spaces eventually combine to become the medullary cavity. 


As the cartilage grows, capillaries penetrate it. This penetration initiates the 
transformation of the perichondrium into the bone-producing periosteum. 
Here, the osteoblasts form a periosteal collar of compact bone around the 
cartilage of the diaphysis. By the second or third month of fetal life, bone 
cell development and ossification ramps up and creates the primary 
ossification center, a region deep in the periosteal collar where ossification 
begins ({link]c). 


While these deep changes are occurring, chondrocytes and cartilage 
continue to grow at the ends of the bone (the future epiphyses), which 
increases the bone’s length at the same time bone is replacing cartilage in 
the diaphyses. By the time the fetal skeleton is fully formed, cartilage only 
remains at the joint surface as articular cartilage and between the diaphysis 
and epiphysis as the epiphyseal plate, the latter of which is responsible for 
the longitudinal growth of bones. After birth, this same sequence of events 
(matrix mineralization, death of chondrocytes, invasion of blood vessels 
from the periosteum, and seeding with osteogenic cells that become 
osteoblasts) occurs in the epiphyseal regions, and each of these centers of 
activity is referred to as a secondary ossification center ((link]e). 


How Bones Grow in Length 


The epiphyseal plate is the area of growth in a long bone. It is a layer of 
hyaline cartilage where ossification occurs in immature bones. On the 
epiphyseal side of the epiphyseal plate, cartilage is formed. On the 
diaphyseal side, cartilage is ossified, and the diaphysis grows in length. The 
epiphyseal plate is composed of four zones of cells and activity ((link]). The 
reserve zone is the region closest to the epiphyseal end of the plate and 
contains small chondrocytes within the matrix. These chondrocytes do not 
participate in bone growth but secure the epiphyseal plate to the osseous 
tissue of the epiphysis. 

Longitudinal Bone Growth 


Changes in 


_— 
Co Q 


Reserve zone fo) (6) Matrix production 


Growth plate zones ere 
= | 


Proliferative zone Mitosis 


_ | Lipids, glycogen, 
and alkaline 
| phosphatase 
accumulate; 
matrix calcifies. 


Maturation and nail 
hypertrophy <P 


Calcified matrix = \— Celldeath 


= 
Primary 
spongiosa 
Secondary 
spongiosa 


Zone of 


ossification 


Metaphysis 


The epiphyseal plate is responsible for 
longitudinal bone growth. 


The proliferative zone is the next layer toward the diaphysis and contains 
stacks of slightly larger chondrocytes. It makes new chondrocytes (via 
mitosis) to replace those that die at the diaphyseal end of the plate. 
Chondrocytes in the next layer, the zone of maturation and hypertrophy, 
are older and larger than those in the proliferative zone. The more mature 
cells are situated closer to the diaphyseal end of the plate. The longitudinal 
growth of bone is a result of cellular division in the proliferative zone and 
the maturation of cells in the zone of maturation and hypertrophy. 


Most of the chondrocytes in the zone of calcified matrix, the zone closest 
to the diaphysis, are dead because the matrix around them has calcified. 
Capillaries and osteoblasts from the diaphysis penetrate this zone, and the 
osteoblasts secrete bone tissue on the remaining calcified cartilage. Thus, 
the zone of calcified matrix connects the epiphyseal plate to the diaphysis. 
A bone grows in length when osseous tissue is added to the diaphysis. 


Bones continue to grow in length until early adulthood. The rate of growth 
is controlled by hormones, which will be discussed later. When the 
chondrocytes in the epiphyseal plate cease their proliferation and bone 
replaces the cartilage, longitudinal growth stops. All that remains of the 
epiphyseal plate is the epiphyseal line ([link]). 

Progression from Epiphyseal Plate to Epiphyseal Line 


Epiphysis ee 


WV Metaphysis 


Diaphysis 


Epiphyseal 
Epiphyseal line 
plate 
(growth 


plate) 


Ee Metaphysis — 
J __ |} —— Eriptysis ——{__U 


Femur 
(thighbone) 


(a) Growing long bone (b) Mature long bone 


As a bone matures, the epiphyseal plate 
progresses to an epiphyseal line. (a) 
Epiphyseal plates are visible in a growing 
bone. (b) Epiphyseal lines are the 
remnants of epiphyseal plates in a mature 
bone. 


How Bones Grow in Diameter 


While bones are increasing in length, they are also increasing in diameter; 
growth in diameter can continue even after longitudinal growth ceases. This 
is called appositional growth. Osteoclasts resorb old bone that lines the 
medullary cavity, while osteoblasts, via intramembranous ossification, 
produce new bone tissue beneath the periosteum. The erosion of old bone 
along the medullary cavity and the deposition of new bone beneath the 
periosteum not only increase the diameter of the diaphysis but also increase 
the diameter of the medullary cavity. This process is called modeling. 


Bone Remodeling 


The process in which matrix is resorbed on one surface of a bone and 
deposited on another is known as bone modeling. Modeling primarily takes 
place during a bone’s growth. However, in adult life, bone undergoes 
remodeling, in which resorption of old or damaged bone takes place on the 
same surface where osteoblasts lay new bone to replace that which is 
resorbed. Injury, exercise, and other activities lead to remodeling. Those 
influences are discussed later in the chapter, but even without injury or 
exercise, about 5 to 10 percent of the skeleton is remodeled annually just by 
destroying old bone and renewing it with fresh bone. 


Note: 

Diseases of the... 

Skeletal System 

Osteogenesis imperfecta (OI) is a genetic disease in which bones do not 
form properly and therefore are fragile and break easily. It is also called 
brittle bone disease. The disease is present from birth and affects a person 
throughout life. 

The genetic mutation that causes OI affects the body’s production of 
collagen, one of the critical components of bone matrix. The severity of the 
disease can range from mild to severe. Those with the most severe forms of 
the disease sustain many more fractures than those with a mild form. 
Frequent and multiple fractures typically lead to bone deformities and short 
stature. Bowing of the long bones and curvature of the spine are also 
common in people afflicted with OI. Curvature of the spine makes 
breathing difficult because the lungs are compressed. 

Because collagen is such an important structural protein in many parts of 
the body, people with OI may also experience fragile skin, weak muscles, 
loose joints, easy bruising, frequent nosebleeds, brittle teeth, blue sclera, 
and hearing loss. There is no known cure for OI. Treatment focuses on 
helping the person retain as much independence as possible while 
minimizing fractures and maximizing mobility. Toward that end, safe 
exercises, like swimming, in which the body is less likely to experience 
collisions or compressive forces, are recommended. Braces to support legs, 
ankles, knees, and wrists are used as needed. Canes, walkers, or 
wheelchairs can also help compensate for weaknesses. 


When bones do break, casts, splints, or wraps are used. In some cases, 
metal rods may be surgically implanted into the long bones of the arms and 
legs. Research is currently being conducted on using bisphosphonates to 
treat OI. Smoking and being overweight are especially risky in people with 
OI, since smoking is known to weaken bones, and extra body weight puts 
additional stress on the bones. 


Note: 
eee 


openstax COLLEGE 


lll 


oF 
1s 


Watch this video to see how a bone grows. 


Chapter Review 


All bone formation is a replacement process. Embryos develop a 
cartilaginous skeleton and various membranes. During development, these 
are replaced by bone during the ossification process. In intramembranous 
ossification, bone develops directly from sheets of mesenchymal connective 
tissue. In endochondral ossification, bone develops by replacing hyaline 
cartilage. Activity in the epiphyseal plate enables bones to grow in length. 
Modeling allows bones to grow in diameter. Remodeling occurs as bone is 
resorbed and replaced by new bone. Osteogenesis imperfecta is a genetic 
disease in which collagen production is altered, resulting in fragile, brittle 
bones. 


Review Questions 


Exercise: 


Problem: Why is cartilage slow to heal? 


a. because it eventually develops into bone 

b. because it is semi-solid and flexible 

c. because it does not have a blood supply 

d. because endochondral ossification replaces all cartilage with bone 


Solution: 


C 


Exercise: 


Problem: Why are osteocytes spread out in bone tissue? 


a. They develop from mesenchymal cells. 

b. They are surrounded by osteoid. 

c. They travel through the capillaries. 

d. Formation of osteoid spreads out the osteoblasts that formed the 
ossification centers. 


Solution: 


D 
Exercise: 


Problem: 
In endochondral ossification, what happens to the chondrocytes? 


a. They develop into osteocytes. 

b. They die in the calcified matrix that surrounds them and form the 
medullary cavity. 

c. They grow and form the periosteum. 

d. They group together to form the primary ossification center. 


Solution: 


B 
Exercise: 


Problem: 


Which of the following bones is (are) formed by intramembranous 
ossification? 


a. the metatarsals 

b. the femur 

c. the ribs 

d. the flat bones of the cranium 


Solution: 
D 
Exercise: 
Problem:Bones grow in length due to activity in the 
a. epiphyseal plate 
b. perichondrium 


Cc. periosteum 
d. medullary cavity 


Solution: 


A 


Exercise: 


Problem:Bones grow in diameter due to bone formation 


a. in the medullary cavity 
b. beneath the periosteum 
c. in the epiphyseal plate 
d. within the metaphysis 


Solution: 


B 
Exercise: 


Problem: 


Which of the following represents the correct sequence of zones in the 
epiphyseal plate? 


a. proliferation, reserved, maturation, calcification 
b. maturation, proliferation, reserved, calcification 
c. calcification, maturation, proliferation, reserved 
d. calcification, reserved, proliferation, maturation 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: 


In what ways do intramembranous and endochondral ossification 
differ? 


Solution: 


In intramembranous ossification, bone develops directly from sheets of 
mesenchymal connective tissue, but in endochondral ossification, bone 
develops by replacing hyaline cartilage. Intramembranous ossification 
is complete by the end of the adolescent growth spurt, while 
endochondral ossification lasts into young adulthood. The flat bones of 
the face, most of the cranial bones, and a good deal of the clavicles 
(collarbones) are formed via intramembranous ossification, while 
bones at the base of the skull and the long bones form via 
endochondral ossification. 


Exercise: 
Problem: 


Considering how a long bone develops, what are the similarities and 
differences between a primary and a secondary ossification center? 


Solution: 


A single primary ossification center is present, during endochondral 
ossification, deep in the periosteal collar. Like the primary ossification 
center, secondary ossification centers are present during endochondral 
ossification, but they form later, and there are two of them, one in each 
epiphysis. 


Glossary 


endochondral ossification 
process in which bone forms by replacing hyaline cartilage 


epiphyseal line 
completely ossified remnant of the epiphyseal plate 


intramembranous ossification 
process by which bone forms directly from mesenchymal tissue 


modeling 


process, during bone growth, by which bone is resorbed on one surface 
of a bone and deposited on another 


ossification 
(also, osteogenesis) bone formation 


ossification center 
cluster of osteoblasts found in the early stages of intramembranous 
ossification 


osteoid 
uncalcified bone matrix secreted by osteoblasts 


perichondrium 
membrane that covers cartilage 


primary ossification center 
region, deep in the periosteal collar, where bone development starts 
during endochondral ossification 


proliferative zone 
region of the epiphyseal plate that makes new chondrocytes to replace 
those that die at the diaphyseal end of the plate and contributes to 
longitudinal growth of the epiphyseal plate 


remodeling 
process by which osteoclasts resorb old or damaged bone at the same 
time as and on the same surface where osteoblasts form new bone to 
replace that which is resorbed 


reserve zone 
region of the epiphyseal plate that anchors the plate to the osseous 
tissue of the epiphysis 


secondary ossification center 
region of bone development in the epiphyses 


zone of calcified matrix 


region of the epiphyseal plate closest to the diaphyseal end; functions 
to connect the epiphyseal plate to the diaphysis 


zone of maturation and hypertrophy 
region of the epiphyseal plate where chondrocytes from the 
proliferative zone grow and mature and contribute to the longitudinal 
growth of the epiphyseal plate 


Fractures: Bone Repair 
By the end of this section, you will be able to: 


e Differentiate among the different types of fractures 
e Describe the steps involved in bone repair 


A fracture is a broken bone. It will heal whether or not a physician resets it 
in its anatomical position. If the bone is not reset correctly, the healing 
process will keep the bone in its deformed position. 


When a broken bone is manipulated and set into its natural position without 
surgery, the procedure is called a closed reduction. Open reduction 
requires surgery to expose the fracture and reset the bone. While some 
fractures can be minor, others are quite severe and result in grave 
complications. For example, a fractured diaphysis of the femur has the 
potential to release fat globules into the bloodstream. These can become 
lodged in the capillary beds of the lungs, leading to respiratory distress and 
if not treated quickly, death. 


Types of Fractures 


Fractures are classified by their complexity, location, and other features 
({link]). [link] outlines common types of fractures. Some fractures may be 
described using more than one term because it may have the features of 
more than one type (e.g., an open transverse fracture). 

Types of Fractures 


Open Transverse 


Compare healthy bone with different types 
of fractures: (a) closed fracture, (b) open 
fracture, (c) transverse fracture, (d) spiral 


fracture, (e) comminuted fracture, (f) 
impacted fracture, (g) greenstick fracture, 
and (h) oblique fracture. 


Types of Fractures 


Type of 
fracture 


Transverse 


Oblique 


Spiral 


Comminuted 


Impacted 


Greenstick 


Open (or 
compound) 


Description 
Occurs straight across the long axis of the bone 
Occurs at an angle that is not 90 degrees 


Bone segments are pulled apart as a result of a 
twisting motion 


Several breaks result in many small pieces between 
two large segments 


One fragment is driven into the other, usually as a 
result of compression 


A partial fracture in which only one side of the 
bone is broken 


A fracture in which at least one end of the broken 
bone tears through the skin; carries a high risk of 
infection 


Types of Fractures 


Type of 

fracture Description 

oe kon A fracture in which the skin remains intact 
simple) 


Bone Repair 


When a bone breaks, blood flows from any vessel torn by the fracture. 
These vessels could be in the periosteum, osteons, and/or medullary cavity. 
The blood begins to clot, and about six to eight hours after the fracture, the 
clotting blood has formed a fracture hematoma ([link]a). The disruption 
of blood flow to the bone results in the death of bone cells around the 
fracture. 

Stages in Fracture Repair 


Hematoma New blood vessels 


cae 
eres 


Bony callus 


Spongy bone 


trabecula of spongy bone 
(b) (c) (d) 


The healing of a bone fracture follows a series of 
progressive steps: (a) A fracture hematoma forms. (b) 
Internal and external calli form. (c) Cartilage of the 
calli is replaced by trabecular bone. (d) Remodeling 
occurs. 


Within about 48 hours after the fracture, chondrocytes from the endosteum 
have created an internal callus (plural = calli) by secreting a 
fibrocartilaginous matrix between the two ends of the broken bone, while 
the periosteal chondrocytes and osteoblasts create an external callus of 
hyaline cartilage and bone, respectively, around the outside of the break 
({link]b). This stabilizes the fracture. 


Over the next several weeks, osteoclasts resorb the dead bone; osteogenic 
cells become active, divide, and differentiate into osteoblasts. The cartilage 
in the calli is replaced by trabecular bone via endochondral ossification 
((link]c). 


Eventually, the internal and external calli unite, compact bone replaces 
spongy bone at the outer margins of the fracture, and healing is complete. A 
slight swelling may remain on the outer surface of the bone, but quite often, 
that region undergoes remodeling ([link]d), and no external evidence of the 
fracture remains. 


Note: 


40 


= 
mess’ OPENStAX COLLEGE” 
— 

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. 


ol r H 


Visit this website to review different types of fractures and then take a 
short self-assessment quiz. 


Chapter Review 


Fractured bones may be repaired by closed reduction or open reduction. 
Fractures are classified by their complexity, location, and other features. 
Common types of fractures are transverse, oblique, spiral, comminuted, 


impacted, greenstick, open (or compound), and closed (or simple). Healing 
of fractures begins with the formation of a hematoma, followed by internal 
and external calli. Osteoclasts resorb dead bone, while osteoblasts create 
new bone that replaces the cartilage in the calli. The calli eventually unite, 
remodeling occurs, and healing is complete. 


Review Questions 


Exercise: 


Problem:A fracture can be both 


a. open and closed 

b. open and transverse 

c. transverse and greenstick 
d. greenstick and comminuted 


Solution: 


B 
Exercise: 
Problem: 


How can a fractured diaphysis release fat globules into the 
bloodstream? 


a. The bone pierces fat stores in the skin. 

b. The yellow marrow in the diaphysis is exposed and damaged. 

c. The injury triggers the body to release fat from healthy bones. 

d. The red marrow in the fractured bone releases fat to heal the 
fracture. 


Solution: 


B 


Exercise: 


Problem:In a compound fracture, 


a. the break occurs at an angle to the bone 

b. the broken bone does not tear the skin 

c. one fragment of broken bone is compressed into the other 
d. broken bone pierces the skin 


Solution: 


D 


Exercise: 


Problem:The internal and external calli are replaced by 


a. hyaline cartilage 
b. trabecular bone 

c. osteogenic cells 
d. osteoclasts 


Solution: 


B 
Exercise: 


Problem: 
The first type of bone to form during fracture repair is bone. 


a. compact 
b. lamellar 


c. spongy 
d. dense 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: 


What is the difference between closed reduction and open reduction? 
In what type of fracture would closed reduction most likely occur? In 
what type of fracture would open reduction most likely occur? 


Solution: 


In closed reduction, the broken ends of a fractured bone can be reset 
without surgery. Open reduction requires surgery to return the broken 
ends of the bone to their correct anatomical position. A partial fracture 
would likely require closed reduction. A compound fracture would 
require open reduction. 


Exercise: 
Problem: 


In terms of origin and composition, what are the differences between 
an internal callus and an external callus? 


Solution: 


The internal callus is produced by cells in the endosteum and is 
composed of a fibrocartilaginous matrix. The external callus is 
produced by cells in the periosteum and consists of hyaline cartilage 
and bone. 


Glossary 


closed reduction 
manual manipulation of a broken bone to set it into its natural position 
without surgery 


external callus 
collar of hyaline cartilage and bone that forms around the outside of a 
fracture 


fracture 
broken bone 


fracture hematoma 
blood clot that forms at the site of a broken bone 


internal callus 
fibrocartilaginous matrix, in the endosteal region, between the two 
ends of a broken bone 


open reduction 
surgical exposure of a bone to reset a fracture 


Divisions of the Skeletal System 
By the end of this section, you will be able to: 


e Discuss the functions of the skeletal system 

e Distinguish between the axial skeleton and appendicular skeleton 
¢ Define the axial skeleton and its components 

¢ Define the appendicular skeleton and its components 


The skeletal system includes all of the bones, cartilages, and ligaments of 
the body that support and give shape to the body and body structures. The 
skeleton consists of the bones of the body. For adults, there are 206 bones 
in the skeleton. Younger individuals have higher numbers of bones because 
some bones fuse together during childhood and adolescence to form an 
adult bone. The primary functions of the skeleton are to provide a rigid, 
internal structure that can support the weight of the body against the force 
of gravity, and to provide a structure upon which muscles can act to 
produce movements of the body. The lower portion of the skeleton is 
specialized for stability during walking or running. In contrast, the upper 
skeleton has greater mobility and ranges of motion, features that allow you 
to lift and carry objects or turn your head and trunk. 


In addition to providing for support and movements of the body, the 
skeleton has protective and storage functions. It protects the internal organs, 
including the brain, spinal cord, heart, lungs, and pelvic organs. The bones 
of the skeleton serve as the primary storage site for important minerals such 
as calcium and phosphate. The bone marrow found within bones stores fat 
and houses the blood-cell producing tissue of the body. 


The skeleton is subdivided into two major divisions—the axial and 
appendicular. 


The Axial Skeleton 


The skeleton is subdivided into two major divisions—the axial and 
appendicular. The axial skeleton forms the vertical, central axis of the body 
and includes all bones of the head, neck, chest, and back ((link]). It serves 
to protect the brain, spinal cord, heart, and lungs. It also serves as the 


attachment site for muscles that move the head, neck, and back, and for 
muscles that act across the shoulder and hip joints to move their 
corresponding limbs. 


The axial skeleton of the adult consists of 80 bones, including the skull, the 
vertebral column, and the thoracic cage. The skull is formed by 22 bones. 
Also associated with the head are an additional seven bones, including the 
hyoid bone and the ear ossicles (three small bones found in each middle 
ear). The vertebral column consists of 24 bones, each called a vertebra, 
plus the sacrum and coccyx. The thoracic cage includes the 12 pairs of 
ribs, and the sternum, the flattened bone of the anterior chest. 

Axial and Appendicular Skeleton 


Skull 


Cranial portion 


6 ) Facial portion ee, 
ey 
ee Pectoral (shoulder) girdle ft} 
= 
ae S 
(FRI mn Clavicle 


E =n) 
SS eS) Scapula 
(ee == | Thoracic cage 
—— Sternum 
a Ly 
‘Z pnt Ribs : 
Vertebral yw ZEN y) _ Uppeeilins = Vertebral 
column Gor . Humerus column 
Pelvic J er. = | one fe. h Pelvic 
girdle WZ | Radius wey, girdle 
(hip bones) | é \ Carpals < OC (hip bones) 
fie oT Met Is foe Oe 
g yi ff iN etacarpals /) N | fiw 
f AY | of \ Phalanges y iy) i 7h 
i 


Lower limb 
———————- Femur 
i G&G EE Patella C) 


Key 
Axial skeleton 


Appendicular 
skeleton 


Tarsals 


\_}— Metatarsals 


Phalanges 


Anterior view Posterior view 


The axial skeleton supports the head, neck, back, and 
chest and thus forms the vertical axis of the body. It 


consists of the skull, vertebral column (including the 
sacrum and coccyx), and the thoracic cage, formed by 
the ribs and sternum. The appendicular skeleton is made 
up of all bones of the upper and lower limbs. 


The Appendicular Skeleton 


The appendicular skeleton includes all bones of the upper and lower 
limbs, plus the bones that attach each limb to the axial skeleton. There are 
126 bones in the appendicular skeleton of an adult. The bones of the 
appendicular skeleton are covered in a separate chapter. 


Chapter Review 


The skeletal system includes all of the bones, cartilages, and ligaments of 
the body. It serves to support the body, protect the brain and other internal 
organs, and provides a rigid structure upon which muscles can pull to 
generate body movements. It also stores fat and the tissue responsible for 
the production of blood cells. The skeleton is subdivided into two parts. The 
axial skeleton forms a vertical axis that includes the head, neck, back, and 
chest. It has 80 bones and consists of the skull, vertebral column, and 
thoracic cage. The adult vertebral column consists of 24 vertebrae plus the 
sacrum and coccyx. The thoracic cage is formed by 12 pairs of ribs and the 
sternum. The appendicular skeleton consists of 126 bones in the adult and 
includes all of the bones of the upper and lower limbs plus the bones that 
anchor each limb to the axial skeleton. 


Review Questions 


Exercise: 


Problem: Which of the following is part of the axial skeleton? 


a. shoulder bones 


b. thigh bone 
c. foot bones 
d. vertebral column 


Solution: 
D 
Exercise: 


Problem: Which of the following is a function of the axial skeleton? 


a. allows for movement of the wrist and hand 

b. protects nerves and blood vessels at the elbow 
c. supports trunk of body 

d. allows for movements of the ankle and foot 


Solution: 
C 
Exercise: 
Problem:The axial skeleton 
a. consists of 126 bones 
b. forms the vertical axis of the body 


c. includes all bones of the body trunk and limbs 
d. includes only the bones of the lower limbs 


Solution: 


B 


Critical Thinking Question 


Exercise: 


Problem: Define the two divisions of the skeleton. 


Solution: 


The axial skeleton forms the vertical axis of the body and includes the 
bones of the head, neck, back, and chest of the body. It consists of 80 

bones that include the skull, vertebral column, and thoracic cage. The 
appendicular skeleton consists of 126 bones and includes all bones of 
the upper and lower limbs. 


Exercise: 


Problem: Discuss the functions of the axial skeleton. 


Solution: 


The axial skeleton supports the head, neck, back, and chest of the body 
and allows for movements of these body regions. It also gives bony 
protections for the brain, spinal cord, heart, and lungs; stores fat and 
minerals; and houses the blood-cell producing tissue. 


Glossary 


appendicular skeleton 
all bones of the upper and lower limbs, plus the girdle bones that 
attach each limb to the axial skeleton 


axial skeleton 
central, vertical axis of the body, including the skull, vertebral column, 
and thoracic cage 


coccyx 
small bone located at inferior end of the adult vertebral column that is 
formed by the fusion of four coccygeal vertebrae; also referred to as 
the “tailbone” 


ear ossicles 
three small bones located in the middle ear cavity that serve to transmit 
sound vibrations to the inner ear 


hyoid bone 
small, U-shaped bone located in upper neck that does not contact any 
other bone 


ribs 
thin, curved bones of the chest wall 


sacrum 
single bone located near the inferior end of the adult vertebral column 
that is formed by the fusion of five sacral vertebrae; forms the 
posterior portion of the pelvis 


skeleton 
bones of the body 


skull 
bony structure that forms the head, face, and jaws, and protects the 
brain; consists of 22 bones 


sternum 
flattened bone located at the center of the anterior chest 


thoracic cage 
consists of 12 pairs of ribs and sternum 


vertebra 
individual bone in the neck and back regions of the vertebral column 


vertebral column 
entire sequence of bones that extend from the skull to the tailbone 


The Skull 
By the end of this section, you will be able to: 


List and identify the bones of the brain case and face 

Locate the major suture lines of the skull and name the bones 
associated with each 

Locate and define the boundaries of the anterior, middle, and posterior 
cranial fossae, the temporal fossa, and infratemporal fossa 

Define the paranasal sinuses and identify the location of each 

Name the bones that make up the walls of the orbit and identify the 
openings associated with the orbit 

Identify the bones and structures that form the nasal septum and nasal 
conchae, and locate the hyoid bone 

Identify the bony openings of the skull 


The cranium (skull) is the skeletal structure of the head that supports the 
face and protects the brain. It is subdivided into the facial bones and the 
brain case, or cranial vault ([link]). The facial bones underlie the facial 
structures, form the nasal cavity, enclose the eyeballs, and support the teeth 
of the upper and lower jaws. The rounded brain case surrounds and protects 
the brain and houses the middle and inner ear structures. 


In the adult, the skull consists of 22 individual bones, 21 of which are 
immobile and united into a single unit. The 22nd bone is the mandible 
(lower jaw), which is the only moveable bone of the skull. 

Parts of the Skull 


Brain case 


Facial bones 


The skull consists of the rounded brain 
case that houses the brain and the facial 
bones that form the upper and lower jaws, 
nose, orbits, and other facial structures. 


Watch this video to view a rotating and exploded skull, with color-coded 
bones. Which bone (yellow) is centrally located and joins with most of the 
other bones of the skull? 


Anterior View of Skull 


The anterior skull consists of the facial bones and provides the bony support 
for the eyes and structures of the face. This view of the skull is dominated 
by the openings of the orbits and the nasal cavity. Also seen are the upper 
and lower jaws, with their respective teeth ([link]). 


The orbit is the bony socket that houses the eyeball and muscles that move 
the eyeball or open the upper eyelid. The upper margin of the anterior orbit 
is the supraorbital margin. Located near the midpoint of the supraorbital 
margin is a small opening called the supraorbital foramen. This provides 
for passage of a sensory nerve to the skin of the forehead. Below the orbit is 
the infraorbital foramen, which is the point of emergence for a sensory 
nerve that supplies the anterior face below the orbit. 

Anterior View of Skull 


Coronal suture 


Glabella Frontal bone 


Parietal bone 
Supraorbital foramen 


Supraorbital margin . 
Orbit 
Sphenoid bone : 
Optic canal 
Temporal bone 
Superior orbital fissure 
Ethmoid bone 
Lacrimal bone 
Nasal bone 


. Inferior orbital fissure 
Palatine bone 


Zygomatic bone 

Nasal septum: 
Perpendicular plate Infraorbital foramen 

of ethmoid bone 


Vomer bone Middle nasal concha 


Inferior nasal concha 


Maxilla 


Alveolar process of maxilla 


Alveolar process 
. / of mandible 
Mental foramen ——————__>) € y 
—_—_ Mandible 


Anterior view 


An anterior view of the skull shows the bones that 
form the forehead, orbits (eye sockets), nasal cavity, 
nasal septum, and upper and lower jaws. 


Inside the nasal area of the skull, the nasal cavity is divided into halves by 
the nasal septum. The upper portion of the nasal septum is formed by the 
perpendicular plate of the ethmoid bone and the lower portion is the 
vomer bone. Each side of the nasal cavity is triangular in shape, with a 
broad inferior space that narrows superiorly. When looking into the nasal 
cavity from the front of the skull, two bony plates are seen projecting from 
each lateral wall. The larger of these is the inferior nasal concha, an 
independent bone of the skull. Located just above the inferior concha is the 
middle nasal concha, which is part of the ethmoid bone. A third bony 
plate, also part of the ethmoid bone, is the superior nasal concha. It is 
much smaller and out of sight, above the middle concha. The superior nasal 
concha is located just lateral to the perpendicular plate, in the upper nasal 
cavity. 


Lateral View of Skull 


A view of the lateral skull is dominated by the large, rounded brain case 
above and the upper and lower jaws with their teeth below ([link]). 
Separating these areas is the bridge of bone called the zygomatic arch. The 
zygomatic arch is the bony arch on the side of skull that spans from the 
area of the cheek to just above the ear canal. It is formed by the junction of 
two bony processes: a short anterior component, the temporal process of 
the zygomatic bone (the cheekbone) and a longer posterior portion, the 
zygomatic process of the temporal bone, extending forward from the 
temporal bone. Thus the temporal process (anteriorly) and the zygomatic 
process (posteriorly) join together, like the two ends of a drawbridge, to 
form the zygomatic arch. One of the major muscles that pulls the mandible 
upward during biting and chewing arises from the zygomatic arch. 


On the lateral side of the brain case, above the level of the zygomatic arch, 
is a Shallow space called the temporal fossa. Below the level of the 
zygomatic arch and deep to the vertical portion of the mandible is another 
space called the infratemporal fossa. Both the temporal fossa and 
infratemporal fossa contain muscles that act on the mandible during 
chewing. 


Lateral View of Skull 


Zygomatic arch 


Coronal suture 


Parietal bone Frontal bone 


Greater wing of 

sphenoid bone 

Si tl 
quamous suture Ethmoid bone 


Temporal bone Lacrimal bone 


Squamous ‘ 
temporal Lacrimal fossa 
Zygomatic process ~L_ 

External acoustic 
meatus 


Lambdoid suture 


Nasal bone 


Zygomatic bone 
Mastoid portion Temporal process 
Styloid process 


Mastoid process 


Maxilla 


Articular tubercle 


Occipital bone Mandibular fossa 


Mandible Mental 


protuberance 
of mandible 


Right lateral view 


The lateral skull shows the large rounded brain case, 
zygomatic arch, and the upper and lower jaws. The 
zygomatic arch is formed jointly by the zygomatic 

process of the temporal bone and the temporal process 
of the zygomatic bone. The shallow space above the 
zygomatic arch is the temporal fossa. The space 
inferior to the zygomatic arch and deep to the 
posterior mandible is the infratemporal fossa. 


Bones of the Brain Case 


The brain case contains and protects the brain. The interior space that is 
almost completely occupied by the brain is called the cranial cavity. This 
cavity is bounded superiorly by the rounded top of the skull, which is called 
the calvaria (skullcap), and the lateral and posterior sides of the skull. The 
bones that form the top and sides of the brain case are usually referred to as 
the “flat” bones of the skull. 


The floor of the brain case is referred to as the base of the skull. This is a 
complex area that varies in depth and has numerous openings for the 
passage of cranial nerves, blood vessels, and the spinal cord. Inside the 
skull, the base is subdivided into three large spaces, called the anterior 
cranial fossa, middle cranial fossa, and posterior cranial fossa (fossa = 
“trench or ditch”) ([link]). From anterior to posterior, the fossae increase in 
depth. The shape and depth of each fossa corresponds to the shape and size 
of the brain region that each houses. The boundaries and openings of the 
cranial fossae (singular = fossa) will be described in a later section. 
Cranial Fossae 


Anterior cranial 
fossa 


Middle cranial 
fossa 


Posterior cranial 


fossa 


Brain within —____. 
cranial cavity ; 


Lateral view 


The bones of the brain case 
surround and protect the brain, 
which occupies the cranial cavity. 
The base of the brain case, which 
forms the floor of cranial cavity, is 
subdivided into the shallow anterior 
cranial fossa, the middle cranial 


fossa, and the deep posterior cranial 
fossa. 


The brain case consists of eight bones. These include the paired parietal and 
temporal bones, plus the unpaired frontal, occipital, sphenoid, and ethmoid 
bones. 


Parietal Bone 


The parietal bone forms most of the upper lateral side of the skull (see 
[link]). These are paired bones, with the right and left parietal bones joining 
together at the top of the skull. Each parietal bone is also bounded 
anteriorly by the frontal bone, inferiorly by the temporal bone, and 
posteriorly by the occipital bone. 


Temporal Bone 


The temporal bone forms the lower lateral side of the skull (see [link]). 
Common wisdom has it that the temporal bone (temporal = “time”’) is so 
named because this area of the head (the temple) is where hair typically first 
turns gray, indicating the passage of time. 


The temporal bone is subdivided into several regions ({link]). The flattened, 
upper portion is the squamous portion of the temporal bone. Below this area 
and projecting anteriorly is the zygomatic process of the temporal bone, 
which forms the posterior portion of the zygomatic arch. Posteriorly is the 
mastoid portion of the temporal bone. Projecting inferiorly from this region 
is a large prominence, the mastoid process, which serves as a muscle 
attachment site. The mastoid process can easily be felt on the side of the 
head just behind your earlobe. On the interior of the skull, the petrous 
portion of each temporal bone forms the prominent, diagonally oriented 
petrous ridge in the floor of the cranial cavity. Located inside each petrous 


ridge are small cavities that house the structures of the middle and inner 
ears. 
Temporal Bone 


External acoustic Squamous 
meatus portion 


Zygomatic 
process 


Articular 


Masicid tubercle 


pomiat Mandibular 


fossa 


Mastoid process Styloid process 


A lateral view of the isolated temporal bone shows 
the squamous, mastoid, and zygomatic portions of 
the temporal bone. 


Important landmarks of the temporal bone, as shown in [link], include the 
following: 


e External acoustic meatus (ear canal)—This is the large opening on 
the lateral side of the skull that is associated with the ear. 

¢ Internal acoustic meatus—This opening is located inside the cranial 
cavity, on the medial side of the petrous ridge. It connects to the 
middle and inner ear cavities of the temporal bone. 

e Mandibular fossa—This is the deep, oval-shaped depression located 
on the external base of the skull, just in front of the external acoustic 
meatus. The mandible (lower jaw) joins with the skull at this site as 
part of the temporomandibular joint, which allows for movements of 
the mandible during opening and closing of the mouth. 


e Articular tubercle—The smooth ridge located immediately anterior 
to the mandibular fossa. Both the articular tubercle and mandibular 
fossa contribute to the temporomandibular joint, the joint that provides 
for movements between the temporal bone of the skull and the 
mandible. 

e Styloid process—Posterior to the mandibular fossa on the external 
base of the skull is an elongated, downward bony projection called the 
styloid process, so named because of its resemblance to a stylus (a pen 
or writing tool). This structure serves as an attachment site for several 
small muscles and for a ligament that supports the hyoid bone of the 
neck. (See also [link].) 

¢ Stylomastoid foramen—This small opening is located between the 
styloid process and mastoid process. This is the point of exit for the 
cranial nerve that supplies the facial muscles. 

¢ Carotid canal—tThe carotid canal is a zig-zag shaped tunnel that 
provides passage through the base of the skull for one of the major 
arteries that supplies the brain. Its entrance is located on the outside 
base of the skull, anteromedial to the styloid process. The canal then 
runs anteromedially within the bony base of the skull, and then turns 
upward to its exit in the floor of the middle cranial cavity, above the 
foramen lacerum. 


External and Internal Views of Base of Skull 


Maxilla: 
Palatine process 


Zygomatic bone 


Palatine bone 
(horizontal plate) 


Zygomatic arch 
Medial and lateral 
pterygoid plates 

Articular tubercle 


Sphenoid bone Foramen ovale 


Foramen spinosum Mandibular fossa 


Foramen lacerum External auditory meatus 


Jugular foramen Mastoid process 


Occipital condyle Styloid process 


Temporal bone Stylomastoid foramen 


Foramen magnum F 
g Entrance to carotid canal 


Occipital bone 


Superior nuchal line External occipital 


protuberance 


(a) Inferior view 


Transverse plane 


Frontal bone 


Ethmoid bone: 
Crista galli 


Superior orbital Cribriform plate 


fissure Sphenoid bone: 


Lesser wing 

Hypophyseal eit eels 
Foramen rotundum turcica 
Foramen lacerum 

and exit of carotid 

canal 


Foramen ovale 
Foramen spinosum 


Internal acoustic meatus 
Temporal bone 


Hypoglossal canal 
Petrous portion 


Foramen magnum (petrous ridge) 


Occipital bone Jugular foramen 


Parietal bone 


(b) Superior view 


(a) The hard palate is formed anteriorly by the palatine 
processes of the maxilla bones and posteriorly by the 
horizontal plate of the palatine bones. (b) The complex floor of 
the cranial cavity is formed by the frontal, ethmoid, sphenoid, 


temporal, and occipital bones. The lesser wing of the sphenoid 
bone separates the anterior and middle cranial fossae. The 
petrous ridge (petrous portion of temporal bone) separates the 
middle and posterior cranial fossae. 


Frontal Bone 


The frontal bone is the single bone that forms the forehead. At its anterior 
midline, between the eyebrows, there is a slight depression called the 
glabella (see [link]). The frontal bone also forms the supraorbital margin of 
the orbit. Near the middle of this margin, is the supraorbital foramen, the 
opening that provides passage for a sensory nerve to the forehead. The 
frontal bone is thickened just above each supraorbital margin, forming 
rounded brow ridges. These are located just behind your eyebrows and vary 
in size among individuals, although they are generally larger in males. 
Inside the cranial cavity, the frontal bone extends posteriorly. This flattened 
region forms both the roof of the orbit below and the floor of the anterior 
cranial cavity above (see [link]b). 


Occipital Bone 


The occipital bone is the single bone that forms the posterior skull and 
posterior base of the cranial cavity ([link]; see also [link]). On its outside 
surface, at the posterior midline, is a small protrusion called the external 
occipital protuberance, which serves as an attachment site for a ligament 
of the posterior neck. Lateral to either side of this bump is a superior 
nuchal line (nuchal = “nape” or “posterior neck”). The nuchal lines 
represent the most superior point at which muscles of the neck attach to the 
skull, with only the scalp covering the skull above these lines. On the base 
of the skull, the occipital bone contains the large opening of the foramen 
magnum, which allows for passage of the spinal cord as it exits the skull. 
On either side of the foramen magnum is an oval-shaped occipital condyle. 


These condyles form joints with the first cervical vertebra and thus support 
the skull on top of the vertebral column. 
Posterior View of Skull 


Parietal bones Sagittal suture 


Lambdoid 
suture 


Occipital bone External 


occipital 
protuberance 


Temporal bone Superior 


nuchal line 


Mastoid process Occipital 


condyle 


Foramen magnum 
Zygomatic 
bone 


Posterior view 


This view of the posterior skull shows 
attachment sites for muscles and joints that 
support the skull. 


Sphenoid Bone 


The sphenoid bone is a single, complex bone of the central skull ([{link]). It 
serves as a “keystone” bone, because it joins with almost every other bone 
of the skull. The sphenoid forms much of the base of the central skull (see 
[link]) and also extends laterally to contribute to the sides of the skull (see 
[link]). Inside the cranial cavity, the right and left lesser wings of the 
sphenoid bone, which resemble the wings of a flying bird, form the lip of a 
prominent ridge that marks the boundary between the anterior and middle 
cranial fossae. The sella turcica (“Turkish saddle”) is located at the midline 
of the middle cranial fossa. This bony region of the sphenoid bone is named 


for its resemblance to the horse saddles used by the Ottoman Turks, with a 
high back and a tall front. The rounded depression in the floor of the sella 
turcica is the hypophyseal (pituitary) fossa, which houses the pea-sized 
pituitary (hypophyseal) gland. The greater wings of the sphenoid bone 
extend laterally to either side away from the sella turcica, where they form 
the anterior floor of the middle cranial fossa. The greater wing is best seen 
on the outside of the lateral skull, where it forms a rectangular area 
immediately anterior to the squamous portion of the temporal bone. 


On the inferior aspect of the skull, each half of the sphenoid bone forms two 
thin, vertically oriented bony plates. These are the medial pterygoid plate 
and lateral pterygoid plate (pterygoid = “wing-shaped”). The right and left 
medial pterygoid plates form the posterior, lateral walls of the nasal cavity. 
The somewhat larger lateral pterygoid plates serve as attachment sites for 
chewing muscles that fill the infratemporal space and act on the mandible. 
Sphenoid Bone 


Superior 
orbital 
fissure 


Foramen 
rotundum 


< 'y Foramen 


ovale 
Ww 
“| Foramen 


spinosum 


Hypophyseal 
fossa of sella 
turcica 


Body of sphenoid 


(a) Superior view 


Body of sphenoid Lesser 
wing 


Superior 
orbital 
fissure 


Pterygoid 
plates 


(b) Posterior view 


Shown in isolation in (a) superior and (b) 
posterior views, the sphenoid bone is a 
single midline bone that forms the 
anterior walls and floor of the middle 
cranial fossa. It has a pair of lesser wings 
and a pair of greater wings. The sella 
turcica surrounds the hypophyseal fossa. 
Projecting downward are the medial and 
lateral pterygoid plates. The sphenoid has 
multiple openings for the passage of 
nerves and blood vessels, including the 
optic canal, superior orbital fissure, 
foramen rotundum, foramen ovale, and 
foramen spinosum. 


Ethmoid Bone 


The ethmoid bone is a single, midline bone that forms the roof and lateral 
walls of the upper nasal cavity, the upper portion of the nasal septum, and 
contributes to the medial wall of the orbit ({link] and [link]). On the interior 
of the skull, the ethmoid also forms a portion of the floor of the anterior 
cranial cavity (see [link |b). 


Within the nasal cavity, the perpendicular plate of the ethmoid bone forms 
the upper portion of the nasal septum. The ethmoid bone also forms the 
lateral walls of the upper nasal cavity. Extending from each lateral wall are 
the superior nasal concha and middle nasal concha, which are thin, curved 
projections that extend into the nasal cavity ({link]). 


In the cranial cavity, the ethmoid bone forms a small area at the midline in 
the floor of the anterior cranial fossa. This region also forms the narrow 
roof of the underlying nasal cavity. This portion of the ethmoid bone 
consists of two parts, the crista galli and cribriform plates. The crista galli 


(“rooster’s comb or crest”) is a small upward bony projection located at the 
midline. It functions as an anterior attachment point for one of the covering 
layers of the brain. To either side of the crista galli is the cribriform plate 
(cribrum = “sieve”), a small, flattened area with numerous small openings 
termed olfactory foramina. Small nerve branches from the olfactory areas of 
the nasal cavity pass through these openings to enter the brain. 


The lateral portions of the ethmoid bone are located between the orbit and 
upper nasal cavity, and thus form the lateral nasal cavity wall and a portion 
of the medial orbit wall. Located inside this portion of the ethmoid bone are 
several small, air-filled spaces that are part of the paranasal sinus system of 
the skull. 

Sagittal Section of Skull 


Sella turcica: 
Hypophyseal fossa 


Parietal bone Crista galli 


Frontal sinus 
Cribriform plate 
Perpendicular plate 


Temporal bone Nasal bone 


F Sphenoid bone 
Internal acoustic 
meatus Sphenoid sinus 


Hypoglossal canal Inferior nasal 


concha 
Occipital bone 


Vomer 
Styloid process 
bs a Maxilla 
Medial and lateral 


pterygoid plates Palatine bone 


Mandibular foramen Mylohyoid line 
Mandible 


Hyoid bone 


This midline view of the sagittally sectioned skull shows the 
nasal septum. 


Ethmoid Bone 


Superior 


Crista galli 
Cribriform plate 


Ethmoid 
air cells 

Superior nasal 

; concha 

Nasal cavity 

Middle nasal concha 
Medial wall 
of orbit Perpendicular plate 


Inferior 


The unpaired ethmoid bone is located at the 
midline within the central skull. It has an upward 
projection, the crista galli, and a downward 
projection, the perpendicular plate, which forms 
the upper nasal septum. The cribriform plates form 
both the roof of the nasal cavity and a portion of 
the anterior cranial fossa floor. The lateral sides of 
the ethmoid bone form the lateral walls of the 
upper nasal cavity, part of the medial orbit wall, 
and give rise to the superior and middle nasal 
conchae. The ethmoid bone also contains the 
ethmoid air cells. 


Lateral Wall of Nasal Cavity 


Ethmoid bone: 
Superior nasal 
concha 
Middle nasal 
concha 
Inferior nasal 
concha 


Sphenoidal 
sinus 


Medial view 


The three nasal conchae are curved bones that 
project from the lateral walls of the nasal cavity. 
The superior nasal concha and middle nasal 
concha are parts of the ethmoid bone. The inferior 
nasal concha is an independent bone of the skull. 


Sutures of the Skull 


A suture is an immobile joint between adjacent bones of the skull. The 
narrow gap between the bones is filled with dense, fibrous connective tissue 
that unites the bones. The long sutures located between the bones of the 
brain case are not straight, but instead follow irregular, tightly twisting 
paths. These twisting lines serve to tightly interlock the adjacent bones, thus 
adding strength to the skull for brain protection. 


The two suture lines seen on the top of the skull are the coronal and sagittal 
sutures. The coronal suture runs from side to side across the skull, within 
the coronal plane of section (see [link]). It joins the frontal bone to the right 
and left parietal bones. The sagittal suture extends posteriorly from the 


coronal suture, running along the midline at the top of the skull in the 
Sagittal plane of section (see [link]). It unites the right and left parietal 
bones. On the posterior skull, the sagittal suture terminates by joining the 
lambdoid suture. The lambdoid suture extends downward and laterally to 
either side away from its junction with the sagittal suture. The lambdoid 
suture joins the occipital bone to the right and left parietal and temporal 
bones. This suture is named for its upside-down "V" shape, which 
resembles the capital letter version of the Greek letter lambda (A). The 
squamous suture is located on the lateral skull. It unites the squamous 
portion of the temporal bone with the parietal bone (see [link]). At the 
intersection of four bones is the pterion, a small, capital-H-shaped suture 
line region that unites the frontal bone, parietal bone, squamous portion of 
the temporal bone, and greater wing of the sphenoid bone. It is the weakest 
part of the skull. The pterion is located approximately two finger widths 
above the zygomatic arch and a thumb’s width posterior to the upward 
portion of the zygomatic bone. 


Note: 

Disorders of the... 

Skeletal System 

Head and traumatic brain injuries are major causes of immediate death and 
disability, with bleeding and infections as possible additional 
complications. According to the Centers for Disease Control and 
Prevention (2010), approximately 30 percent of all injury-related deaths in 
the United States are caused by head injuries. The majority of head injuries 
involve falls. They are most common among young children (ages 0-4 
years), adolescents (15-19 years), and the elderly (over 65 years). 
Additional causes vary, but prominent among these are automobile and 
motorcycle accidents. 

Strong blows to the brain-case portion of the skull can produce fractures. 
These may result in bleeding inside the skull with subsequent injury to the 
brain. The most common is a linear skull fracture, in which fracture lines 
radiate from the point of impact. Other fracture types include a 
comminuted fracture, in which the bone is broken into several pieces at the 
point of impact, or a depressed fracture, in which the fractured bone is 


pushed inward. In a contrecoup (counterblow) fracture, the bone at the 
point of impact is not broken, but instead a fracture occurs on the opposite 
side of the skull. Fractures of the occipital bone at the base of the skull can 
occur in this manner, producing a basilar fracture that can damage the 
artery that passes through the carotid canal. 

A blow to the lateral side of the head may fracture the bones of the pterion. 
The pterion is an important clinical landmark because located immediately 
deep to it on the inside of the skull is a major branch of an artery that 
supplies the skull and covering layers of the brain. A strong blow to this 
region can fracture the bones around the pterion. If the underlying artery is 
damaged, bleeding can cause the formation of a hematoma (collection of 
blood) between the brain and interior of the skull. As blood accumulates, it 
will put pressure on the brain. Symptoms associated with a hematoma may 
not be apparent immediately following the injury, but if untreated, blood 
accumulation will exert increasing pressure on the brain and can result in 
death within a few hours. 


Note: 

[= [=] 
‘all 

“eat a ee 

5 openstax COLLEGE” 


View this animation to see how a blow to the head may produce a 
contrecoup (counterblow) fracture of the basilar portion of the occipital 
bone on the base of the skull. Why may a basilar fracture be life 
threatening? 


Facial Bones of the Skull 


The facial bones of the skull form the upper and lower jaws, the nose, nasal 
cavity and nasal septum, and the orbit. The facial bones include 14 bones, 
with six paired bones and two unpaired bones. The paired bones are the 
maxilla, palatine, zygomatic, nasal, lacrimal, and inferior nasal conchae 
bones. The unpaired bones are the vomer and mandible bones. Although 
classified with the brain-case bones, the ethmoid bone also contributes to 
the nasal septum and the walls of the nasal cavity and orbit. 


Maxillary Bone 


The maxillary bone, often referred to simply as the maxilla (plural = 
maxillae), is one of a pair that together form the upper jaw, much of the 
hard palate, the medial floor of the orbit, and the lateral base of the nose 
(see [link]). The curved, inferior margin of the maxillary bone that forms 
the upper jaw and contains the upper teeth is the alveolar process of the 
maxilla ({link]). Each tooth is anchored into a deep socket called an 
alveolus. On the anterior maxilla, just below the orbit, is the infraorbital 
foramen. This is the point of exit for a sensory nerve that supplies the nose, 
upper lip, and anterior cheek. On the inferior skull, the palatine process 
from each maxillary bone can be seen joining together at the midline to 
form the anterior three-quarters of the hard palate (see [link]a). The hard 
palate is the bony plate that forms the roof of the mouth and floor of the 
nasal cavity, separating the oral and nasal cavities. 

Maxillary Bone 


Articulates with 
frontal bone 


Infraorbital 
foramen 


Alveolar 
process 


Right lateral view 


The maxillary bone forms the upper jaw and 
supports the upper teeth. Each maxilla also 
forms the lateral floor of each orbit and the 

majority of the hard palate. 


Palatine Bone 


The palatine bone is one of a pair of irregularly shaped bones that 
contribute small areas to the lateral walls of the nasal cavity and the medial 
wall of each orbit. The largest region of each of the palatine bone is the 
horizontal plate. The plates from the right and left palatine bones join 
together at the midline to form the posterior quarter of the hard palate (see 
[link]a). Thus, the palatine bones are best seen in an inferior view of the 
skull and hard palate. 


Note: 
Homeostatic Imbalances 


Cleft Lip and Cleft Palate 

During embryonic development, the right and left maxilla bones come 
together at the midline to form the upper jaw. At the same time, the muscle 
and skin overlying these bones join together to form the upper lip. Inside 
the mouth, the palatine processes of the maxilla bones, along with the 
horizontal plates of the right and left palatine bones, join together to form 
the hard palate. If an error occurs in these developmental processes, a birth 
defect of cleft lip or cleft palate may result. 

Cleft lip is a common development defect that affects approximately 
1:1000 births, most of which are male. This defect involves a partial or 
complete failure of the right and left portions of the upper lip to fuse 
together, leaving a cleft (gap). 

A more severe developmental defect is cleft palate, which affects the hard 
palate. The hard palate is the bony structure that separates the nasal cavity 
from the oral cavity. It is formed during embryonic development by the 
midline fusion of the horizontal plates from the right and left palatine 
bones and the palatine processes of the maxilla bones. Cleft palate affects 
approximately 1:2500 births and is more common in females. It results 
from a failure of the two halves of the hard palate to completely come 
together and fuse at the midline, thus leaving a gap between them. This gap 
allows for communication between the nasal and oral cavities. In severe 
cases, the bony gap continues into the anterior upper jaw where the 
alveolar processes of the maxilla bones also do not properly join together 
above the front teeth. If this occurs, a cleft lip will also be seen. Because of 
the communication between the oral and nasal cavities, a cleft palate 
makes it very difficult for an infant to generate the suckling needed for 
nursing, thus leaving the infant at risk for malnutrition. Surgical repair is 
required to correct cleft palate defects. 


Zygomatic Bone 


The zygomatic bone is also known as the cheekbone. Each of the paired 
zygomatic bones forms much of the lateral wall of the orbit and the lateral- 
inferior margins of the anterior orbital opening (see [link]). The short 


temporal process of the zygomatic bone projects posteriorly, where it forms 
the anterior portion of the zygomatic arch (see [link]). 


Nasal Bone 


The nasal bone is one of two small bones that articulate (join) with each 
other to form the bony base (bridge) of the nose. They also support the 
cartilages that form the lateral walls of the nose (see [link]). These are the 
bones that are damaged when the nose is broken. 


Lacrimal Bone 


Each lacrimal bone is a small, rectangular bone that forms the anterior, 
medial wall of the orbit (see [link] and [link]). The anterior portion of the 
lacrimal bone forms a shallow depression called the lacrimal fossa, and 
extending inferiorly from this is the nasolacrimal canal. The lacrimal fluid 
(tears of the eye), which serves to maintain the moist surface of the eye, 
drains at the medial corner of the eye into the nasolacrimal canal. This duct 
then extends downward to open into the nasal cavity, behind the inferior 
nasal concha. In the nasal cavity, the lacrimal fluid normally drains 
posteriorly, but with an increased flow of tears due to crying or eye 
irritation, some fluid will also drain anteriorly, thus causing a runny nose. 


Inferior Nasal Conchae 


The right and left inferior nasal conchae form a curved bony plate that 
projects into the nasal cavity space from the lower lateral wall (see [link]). 
The inferior concha is the largest of the nasal conchae and can easily be 
seen when looking into the anterior opening of the nasal cavity. 


Vomer Bone 


The unpaired vomer bone, often referred to simply as the vomer, is 
triangular-shaped and forms the posterior-inferior part of the nasal septum 
(see [link]). The vomer is best seen when looking from behind into the 
posterior openings of the nasal cavity (see [link]a). In this view, the vomer 
is seen to form the entire height of the nasal septum. A much smaller 
portion of the vomer can also be seen when looking into the anterior 
opening of the nasal cavity. 


Mandible 


The mandible forms the lower jaw and is the only moveable bone of the 
skull. At the time of birth, the mandible consists of paired right and left 
bones, but these fuse together during the first year to form the single U- 
shaped mandible of the adult skull. Each side of the mandible consists of a 
horizontal body and posteriorly, a vertically oriented ramus of the 
mandible (ramus = “branch’). The outside margin of the mandible, where 
the body and ramus come together is called the angle of the mandible 
({link]). 


The ramus on each side of the mandible has two upward-going bony 
projections. The more anterior projection is the flattened coronoid process 
of the mandible, which provides attachment for one of the biting muscles. 
The posterior projection is the condylar process of the mandible, which is 
topped by the oval-shaped condyle. The condyle of the mandible articulates 
(joins) with the mandibular fossa and articular tubercle of the temporal 
bone. Together these articulations form the temporomandibular joint, which 
allows for opening and closing of the mouth (see [link]). The broad U- 
shaped curve located between the coronoid and condylar processes is the 
mandibular notch. 


Important landmarks for the mandible include the following: 


e Alveolar process of the mandible—This is the upper border of the 
mandibular body and serves to anchor the lower teeth. 

¢ Mental protuberance—The forward projection from the inferior 
margin of the anterior mandible that forms the chin (mental = “chin’”). 


¢ Mental foramen—The opening located on each side of the anterior- 
lateral mandible, which is the exit site for a sensory nerve that supplies 
the chin. 

e Mylohyoid line—This bony ridge extends along the inner aspect of 
the mandibular body (see [link]). The muscle that forms the floor of 
the oral cavity attaches to the mylohyoid lines on both sides of the 
mandible. 

e Mandibular foramen—This opening is located on the medial side of 
the ramus of the mandible. The opening leads into a tunnel that runs 
down the length of the mandibular body. The sensory nerve and blood 
vessels that supply the lower teeth enter the mandibular foramen and 
then follow this tunnel. Thus, to numb the lower teeth prior to dental 
work, the dentist must inject anesthesia into the lateral wall of the oral 
cavity at a point prior to where this sensory nerve enters the 
mandibular foramen. 

¢ Lingula—tThis small flap of bone is named for its shape (lingula = 
“little tongue”). It is located immediately next to the mandibular 
foramen, on the medial side of the ramus. A ligament that anchors the 
mandible during opening and closing of the mouth extends down from 
the base of the skull and attaches to the lingula. 


Isolated Mandible 


Condylar process ; 
Coronoid process 


Lingula 


Mandibular 
notch 


Mandibular 
condyle 


Mandibular foramen 
Mylohyoid line 
Alveolar process 


Ramus of 
mandible 


Mental 
protuberance 


Mental foramen 


Mandibular 
angle 


Body of mandible 


Right lateral view 


The mandible is the only moveable bone of the 
skull. 


The Orbit 


The orbit is the bony socket that houses the eyeball and contains the 
muscles that move the eyeball or open the upper eyelid. Each orbit is cone- 
shaped, with a narrow posterior region that widens toward the large anterior 
opening. To help protect the eye, the bony margins of the anterior opening 
are thickened and somewhat constricted. The medial walls of the two orbits 
are parallel to each other but each lateral wall diverges away from the 
midline at a 45° angle. This divergence provides greater lateral peripheral 
vision. 


The walls of each orbit include contributions from seven skull bones 
({link]). The frontal bone forms the roof and the zygomatic bone forms the 
lateral wall and lateral floor. The medial floor is primarily formed by the 
maxilla, with a small contribution from the palatine bone. The ethmoid 
bone and lacrimal bone make up much of the medial wall and the sphenoid 
bone forms the posterior orbit. 


At the posterior apex of the orbit is the opening of the optic canal, which 
allows for passage of the optic nerve from the retina to the brain. Lateral to 
this is the elongated and irregularly shaped superior orbital fissure, which 
provides passage for the artery that supplies the eyeball, sensory nerves, and 
the nerves that supply the muscles involved in eye movements. 

Bones of the Orbit 


Frontal bone Supraorbital foramen 


Supraorbital margin 


Sphenoid bone Nasal bone 


Lacrimal bone 
Ethmoid bone 
Lacrimal fossa 


Optic canal 


Superior orbital fissure 


Zygomatic bone Palatine bone 


Infraorbital foramen 


Seven skull bones contribute to the walls of the orbit. Opening 
into the posterior orbit from the cranial cavity are the optic 
canal and superior orbital fissure. 


The Nasal Septum and Nasal Conchae 


The nasal septum consists of both bone and cartilage components ([Link]; 
see also [link]). The upper portion of the septum is formed by the 
perpendicular plate of the ethmoid bone. The lower and posterior parts of 
the septum are formed by the triangular-shaped vomer bone. In an anterior 
view of the skull, the perpendicular plate of the ethmoid bone is easily seen 
inside the nasal opening as the upper nasal septum, but only a small portion 
of the vomer is seen as the inferior septum. A better view of the vomer bone 
is seen when looking into the posterior nasal cavity with an inferior view of 
the skull, where the vomer forms the full height of the nasal septum. The 
anterior nasal septum is formed by the septal cartilage, a flexible plate that 
fills in the gap between the perpendicular plate of the ethmoid and vomer 
bones. This cartilage also extends outward into the nose where it separates 
the right and left nostrils. The septal cartilage is not found in the dry skull. 


Attached to the lateral wall on each side of the nasal cavity are the superior, 
middle, and inferior nasal conchae (singular = concha), which are named 


for their positions (see [link]). These are bony plates that curve downward 
as they project into the space of the nasal cavity. They serve to swirl the 
incoming air, which helps to warm and moisturize it before the air moves 
into the delicate air sacs of the lungs. This also allows mucus, secreted by 
the tissue lining the nasal cavity, to trap incoming dust, pollen, bacteria, and 
viruses. The largest of the conchae is the inferior nasal concha, which is an 
independent bone of the skull. The middle concha and the superior conchae, 
which is the smallest, are both formed by the ethmoid bone. When looking 
into the anterior nasal opening of the skull, only the inferior and middle 
conchae can be seen. The small superior nasal concha is well hidden above 
and behind the middle concha. 

Nasal Septum 


Frontal bone 


Crista galli 


Frontal sinus 


Sphenoid sinus Nasal bone 


Nasal septum: 
Perpendicular plate 
of ethmoid bone 


Vomer bone 


Septal cartilage 


Sphenoid 
bone 

Qy Palatine process 
Horizontal plate — of maxilla 
of palatine bone 


Sagittal section 


The nasal septum is formed by the perpendicular plate of the 
ethmoid bone and the vomer bone. The septal cartilage fills the 
gap between these bones and extends into the nose. 


Cranial Fossae 


Inside the skull, the floor of the cranial cavity is subdivided into three 
cranial fossae (spaces), which increase in depth from anterior to posterior 
(see [link], [link]b, and [link]). Since the brain occupies these areas, the 
shape of each conforms to the shape of the brain regions that it contains. 
Each cranial fossa has anterior and posterior boundaries and is divided at 
the midline into right and left areas by a significant bony structure or 
opening. 


Anterior Cranial Fossa 


The anterior cranial fossa is the most anterior and the shallowest of the 
three cranial fossae. It overlies the orbits and contains the frontal lobes of 
the brain. Anteriorly, the anterior fossa is bounded by the frontal bone, 
which also forms the majority of the floor for this space. The lesser wings 
of the sphenoid bone form the prominent ledge that marks the boundary 
between the anterior and middle cranial fossae. Located in the floor of the 
anterior cranial fossa at the midline is a portion of the ethmoid bone, 
consisting of the upward projecting crista galli and to either side of this, the 
cribriform plates. 


Middle Cranial Fossa 


The middle cranial fossa is deeper and situated posterior to the anterior 
fossa. It extends from the lesser wings of the sphenoid bone anteriorly, to 
the petrous ridges (petrous portion of the temporal bones) posteriorly. The 
large, diagonally positioned petrous ridges give the middle cranial fossa a 
butterfly shape, making it narrow at the midline and broad laterally. The 
temporal lobes of the brain occupy this fossa. The middle cranial fossa is 
divided at the midline by the upward bony prominence of the sella turcica, a 
part of the sphenoid bone. The middle cranial fossa has several openings for 
the passage of blood vessels and cranial nerves (see [link]). 


Openings in the middle cranial fossa are as follows: 


Optic canal—This opening is located at the anterior lateral comer of 
the sella turcica. It provides for passage of the optic nerve into the 
orbit. 

Superior orbital fissure—This large, irregular opening into the 
posterior orbit is located on the anterior wall of the middle cranial 
fossa, lateral to the optic canal and under the projecting margin of the 
lesser wing of the sphenoid bone. Nerves to the eyeball and associated 
muscles, and sensory nerves to the forehead pass through this opening. 
Foramen rotundum—This rounded opening (rotundum = “round”) is 
located in the floor of the middle cranial fossa, just inferior to the 
superior orbital fissure. It is the exit point for a major sensory nerve 
that supplies the cheek, nose, and upper teeth. 

Foramen ovale of the middle cranial fossa—This large, oval-shaped 
opening in the floor of the middle cranial fossa provides passage for a 
major sensory nerve to the lateral head, cheek, chin, and lower teeth. 
Foramen spinosum—This small opening, located posterior-lateral to 
the foramen ovale, is the entry point for an important artery that 
supplies the covering layers surrounding the brain. The branching 
pattern of this artery forms readily visible grooves on the internal 
surface of the skull and these grooves can be traced back to their origin 
at the foramen spinosum. 

Carotid canal—This is the zig-zag passageway through which a 
major artery to the brain enters the skull. The entrance to the carotid 
canal is located on the inferior aspect of the skull, anteromedial to the 
styloid process (see [link]a). From here, the canal runs anteromedially 
within the bony base of the skull. Just above the foramen lacerum, the 
carotid canal opens into the middle cranial cavity, near the posterior- 
lateral base of the sella turcica. 

Foramen lacerum—This irregular opening is located in the base of 
the skull, immediately inferior to the exit of the carotid canal. This 
opening is an artifact of the dry skull, because in life it is completely 
filled with cartilage. All the openings of the skull that provide for 
passage of nerves or blood vessels have smooth margins; the word 
lacerum (“ragged” or “torn’”) tells us that this opening has ragged 
edges and thus nothing passes through it. 


Posterior Cranial Fossa 


The posterior cranial fossa is the most posterior and deepest portion of the 
cranial cavity. It contains the cerebellum of the brain. The posterior fossa is 
bounded anteriorly by the petrous ridges, while the occipital bone forms the 
floor and posterior wall. It is divided at the midline by the large foramen 
magnum (“great aperture”), the opening that provides for passage of the 
spinal cord. 


Located on the medial wall of the petrous ridge in the posterior cranial fossa 
is the internal acoustic meatus (see [link]). This opening provides for 
passage of the nerve from the hearing and equilibrium organs of the inner 
ear, and the nerve that supplies the muscles of the face. Located at the 
anterior-lateral margin of the foramen magnum is the hypoglossal canal. 
These emerge on the inferior aspect of the skull at the base of the occipital 
condyle and provide passage for an important nerve to the tongue. 


Immediately inferior to the internal acoustic meatus is the large, irregularly 
shaped jugular foramen (see [link]a). Several cranial nerves from the brain 
exit the skull via this opening. It is also the exit point through the base of 
the skull for all the venous return blood leaving the brain. The venous 
structures that carry blood inside the skull form large, curved grooves on 
the inner walls of the posterior cranial fossa, which terminate at each 
jugular foramen. 


Paranasal Sinuses 


The paranasal sinuses are hollow, air-filled spaces located within certain 
bones of the skull ([link]). All of the sinuses communicate with the nasal 
cavity (paranasal = “next to nasal cavity”) and are lined with nasal mucosa. 
They serve to reduce bone mass and thus lighten the skull, and they also 
add resonance to the voice. This second feature is most obvious when you 
have a cold or sinus congestion. These produce swelling of the mucosa and 
excess mucus production, which can obstruct the narrow passageways 
between the sinuses and the nasal cavity, causing your voice to sound 
different to yourself and others. This blockage can also allow the sinuses to 
fill with fluid, with the resulting pressure producing pain and discomfort. 


The paranasal sinuses are named for the skull bone that each occupies. The 
frontal sinus is located just above the eyebrows, within the frontal bone 
(see [link]). This irregular space may be divided at the midline into bilateral 
spaces, or these may be fused into a single sinus space. The frontal sinus is 
the most anterior of the paranasal sinuses. The largest sinus is the maxillary 
sinus. These are paired and located within the right and left maxillary 
bones, where they occupy the area just below the orbits. The maxillary 
sinuses are most commonly involved during sinus infections. Because their 
connection to the nasal cavity is located high on their medial wall, they are 
difficult to drain. The sphenoid sinus is a single, midline sinus. It is located 
within the body of the sphenoid bone, just anterior and inferior to the sella 
turcica, thus making it the most posterior of the paranasal sinuses. The 
lateral aspects of the ethmoid bone contain multiple small spaces separated 
by very thin bony walls. Each of these spaces is called an ethmoid air cell. 
These are located on both sides of the ethmoid bone, between the upper 
nasal cavity and medial orbit, just behind the superior nasal conchae. 
Paranasal Sinuses 


Anterior Lateral 


The paranasal sinuses are hollow, air-filled spaces 
named for the skull bone that each occupies. The most 
anterior is the frontal sinus, located in the frontal bone 

above the eyebrows. The largest are the maxillary 

sinuses, located in the right and left maxillary bones 


below the orbits. The most posterior is the sphenoid 
sinus, located in the body of the sphenoid bone, under 
the sella turcica. The ethmoid air cells are multiple 
small spaces located in the right and left sides of the 
ethmoid bone, between the medial wall of the orbit 
and lateral wall of the upper nasal cavity. 


Hyoid Bone 


The hyoid bone is an independent bone that does not contact any other bone 
and thus is not part of the skull ({link]). It is a small U-shaped bone located 
in the upper neck near the level of the inferior mandible, with the tips of the 
“U” pointing posteriorly. The hyoid serves as the base for the tongue above, 
and is attached to the larynx below and the pharynx posteriorly. The hyoid 
is held in position by a series of small muscles that attach to it either from 
above or below. These muscles act to move the hyoid up/down or 
forward/back. Movements of the hyoid are coordinated with movements of 
the tongue, larynx, and pharynx during swallowing and speaking. 

Hyoid Bone 


Mandible 


Hyoid bone 


Larynx 


Greater horn 


Lesser horn 


Body 


Greater horn 


Lesser horn 


Right lateral view 


The hyoid bone is located in the 
upper neck and does not join 
with any other bone. It provides 
attachments for muscles that act 
on the tongue, larynx, and 
pharynx. 


Chapter Review 


The skull consists of the brain case and the facial bones. The brain case 
surrounds and protects the brain, which occupies the cranial cavity inside 
the skull. It consists of the rounded calvaria and a complex base. The brain 
case is formed by eight bones, the paired parietal and temporal bones plus 


the unpaired frontal, occipital, sphenoid, and ethmoid bones. The narrow 
gap between the bones is filled with dense, fibrous connective tissue that 
unites the bones. The sagittal suture joins the right and left parietal bones. 
The coronal suture joins the parietal bones to the frontal bone, the lamboid 
suture joins them to the occipital bone, and the squamous suture joins them 
to the temporal bone. 


The facial bones support the facial structures and form the upper and lower 
jaws. These consist of 14 bones, with the paired maxillary, palatine, 
zygomatic, nasal, lacrimal, and inferior conchae bones and the unpaired 
vomer and mandible bones. The ethmoid bone also contributes to the 
formation of facial structures. The maxilla forms the upper jaw and the 
mandible forms the lower jaw. The maxilla also forms the larger anterior 
portion of the hard palate, which is completed by the smaller palatine bones 
that form the posterior portion of the hard palate. 


The floor of the cranial cavity increases in depth from front to back and is 
divided into three cranial fossae. The anterior cranial fossa is located 
between the frontal bone and lesser wing of the sphenoid bone. A small 
area of the ethmoid bone, consisting of the crista galli and cribriform plates, 
is located at the midline of this fossa. The middle cranial fossa extends from 
the lesser wing of the sphenoid bone to the petrous ridge (petrous portion of 
temporal bone). The right and left sides are separated at the midline by the 
sella turcica, which surrounds the shallow hypophyseal fossa. Openings 
through the skull in the floor of the middle fossa include the optic canal and 
superior orbital fissure, which open into the posterior orbit, the foramen 
rotundum, foramen ovale, and foramen spinosum, and the exit of the carotid 
canal with its underlying foramen lacerum. The deep posterior cranial fossa 
extends from the petrous ridge to the occipital bone. Openings here include 
the large foramen magnum, plus the internal acoustic meatus, jugular 
foramina, and hypoglossal canals. Additional openings located on the 
external base of the skull include the stylomastoid foramen and the entrance 
to the carotid canal. 


The anterior skull has the orbits that house the eyeballs and associated 
muscles. The walls of the orbit are formed by contributions from seven 
bones: the frontal, zygomatic, maxillary, palatine, ethmoid, lacrimal, and 


sphenoid. Located at the superior margin of the orbit is the supraorbital 
foramen, and below the orbit is the infraorbital foramen. The mandible has 
two openings, the mandibular foramen on its inner surface and the mental 
foramen on its external surface near the chin. The nasal conchae are bony 
projections from the lateral walls of the nasal cavity. The large inferior 
nasal concha is an independent bone, while the middle and superior 
conchae are parts of the ethmoid bone. The nasal septum is formed by the 
perpendicular plate of the ethmoid bone, the vomer bone, and the septal 
cartilage. The paranasal sinuses are air-filled spaces located within the 
frontal, maxillary, sphenoid, and ethmoid bones. 


On the lateral skull, the zygomatic arch consists of two parts, the temporal 
process of the zygomatic bone anteriorly and the zygomatic process of the 
temporal bone posteriorly. The temporal fossa is the shallow space located 
on the lateral skull above the level of the zygomatic arch. The infratemporal 
fossa is located below the zygomatic arch and deep to the ramus of the 
mandible. 


The hyoid bone is located in the upper neck and does not join with any 
other bone. It is held in position by muscles and serves to support the 
tongue above, the larynx below, and the pharynx posteriorly. 


Interactive Link Questions 


Exercise: 
Problem: 
Watch this video to view a rotating and exploded skull with color- 


coded bones. Which bone (yellow) is centrally located and joins with 
most of the other bones of the skull? 


Solution: 


The sphenoid bone joins with most other bones of the skull. It is 
centrally located, where it forms portions of the rounded brain case 
and cranial base. 


Exercise: 


Problem: 


View this animation to see how a blow to the head may produce a 
contrecoup (counterblow) fracture of the basilar portion of the 
occipital bone on the base of the skull. Why may a basilar fracture be 
life threatening? 


Solution: 


A basilar fracture may damage an artery entering the skull, causing 
bleeding in the brain. 


Review Questions 


Exercise: 


Problem: Which of the following is a bone of the brain case? 


a. parietal bone 
b. zygomatic bone 
c. maxillary bone 
d. lacrimal bone 


Solution: 


A 


Exercise: 


Problem:The lambdoid suture joins the parietal bone to the 


a. frontal bone 

b. occipital bone 

c. other parietal bone 
d. temporal bone 


Solution: 


B 


Exercise: 


Problem:The middle cranial fossa 


a. is bounded anteriorly by the petrous ridge 

b. is bounded posteriorly by the lesser wing of the sphenoid bone 

c. is divided at the midline by a small area of the ethmoid bone 

d. has the foramen rotundum, foramen ovale, and foramen spinosum 


Solution: 


D 


Exercise: 


Problem:The paranasal sinuses are 


a. air-filled spaces found within the frontal, maxilla, sphenoid, and 
ethmoid bones only 

b. air-filled spaces found within all bones of the skull 

c. not connected to the nasal cavity 

d. divided at the midline by the nasal septum 


Solution: 


A 


Exercise: 


Problem: Parts of the sphenoid bone include the 


a. sella turcica 

b. squamous portion 
c. glabella 

d. zygomatic process 


Solution: 
A 
Exercise: 
Problem:The bony openings of the skull include the 


a. carotid canal, which is located in the anterior cranial fossa 

b. superior orbital fissure, which is located at the superior margin of 
the anterior orbit 

c. mental foramen, which is located just below the orbit 

d. hypoglossal canal, which is located in the posterior cranial fossa 


Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: 


Define and list the bones that form the brain case or support the facial 
structures. 


Solution: 


The brain case is that portion of the skull that surrounds and protects 
the brain. It is subdivided into the rounded top of the skull, called the 


calvaria, and the base of the skull. There are eight bones that form the 
brain case. These are the paired parietal and temporal bones, plus the 
unpaired frontal, occipital, sphenoid, and ethmoid bones. The facial 
bones support the facial structures, and form the upper and lower jaws, 
nasal cavity, nasal septum, and orbit. There are 14 facial bones. These 
are the paired maxillary, palatine, zygomatic, nasal, lacrimal, and 
inferior nasal conchae bones, and the unpaired vomer and mandible 
bones. 


Exercise: 


Problem: 


Identify the major sutures of the skull, their locations, and the bones 
united by each. 


Solution: 


The coronal suture passes across the top of the anterior skull. It unites 
the frontal bone anteriorly with the right and left parietal bones. The 
sagittal suture runs at the midline on the top of the skull. It unites the 
right and left parietal bones with each other. The squamous suture is a 
curved suture located on the lateral side of the skull. It unites the 
squamous portion of the temporal bone to the parietal bone. The 
lambdoid suture is located on the posterior skull and has an inverted V- 
shape. It unites the occipital bone with the right and left parietal bones. 


Exercise: 
Problem: 
Describe the anterior, middle, and posterior cranial fossae and their 


boundaries, and give the midline structure that divides each into right 
and left areas. 


Solution: 


The anterior cranial fossa is the shallowest of the three cranial fossae. 
It extends from the frontal bone anteriorly to the lesser wing of the 
sphenoid bone posteriorly. It is divided at the midline by the crista galli 


and cribriform plates of the ethmoid bone. The middle cranial fossa is 
located in the central skull, and is deeper than the anterior fossa. The 
middle fossa extends from the lesser wing of the sphenoid bone 
anteriorly to the petrous ridge posteriorly. It is divided at the midline 
by the sella turcica. The posterior cranial fossa is the deepest fossa. It 
extends from the petrous ridge anteriorly to the occipital bone 
posteriorly. The large foramen magnum is located at the midline of the 
posterior fossa. 


Exercise: 
Problem: 
Describe the parts of the nasal septum in both the dry and living skull. 
Solution: 
There are two bony parts of the nasal septum in the dry skull. The 
perpendicular plate of the ethmoid bone forms the superior part of the 
septum. The vomer bone forms the inferior and posterior parts of the 
septum. In the living skull, the septal cartilage completes the septum 
by filling in the anterior area between the bony components and 
extending outward into the nose. 


References 


Centers for Disease Control and Prevention (US). Injury prevention and 
control: traumatic brain injury [Internet]. Atlanta, GA; [cited 2013 Mar 18]. 


Glossary 


alveolar process of the mandible 
upper border of mandibular body that contains the lower teeth 


alveolar process of the maxilla 


curved, inferior margin of the maxilla that supports and anchors the 
upper teeth 


angle of the mandible 
rounded corner located at outside margin of the body and ramus 
junction 


anterior cranial fossa 
shallowest and most anterior cranial fossa of the cranial base that 
extends from the frontal bone to the lesser wing of the sphenoid bone 


articular tubercle 
smooth ridge located on the inferior skull, immediately anterior to the 
mandibular fossa 


brain case 
portion of the skull that contains and protects the brain, consisting of 
the eight bones that form the cranial base and rounded upper skull 


calvaria 
(also, skullcap) rounded top of the skull 


carotid canal 
zig-zag tunnel providing passage through the base of the skull for the 
internal carotid artery to the brain; begins anteromedial to the styloid 
process and terminates in the middle cranial cavity, near the posterior- 
lateral base of the sella turcica 


condylar process of the mandible 
thickened upward projection from posterior margin of mandibular 
ramus 


condyle 
oval-shaped process located at the top of the condylar process of the 
mandible 


coronal suture 


joint that unites the frontal bone to the right and left parietal bones 
across the top of the skull 


coronoid process of the mandible 
flattened upward projection from the anterior margin of the mandibular 
ramus 


cranial cavity 
interior space of the skull that houses the brain 


cranium 
skull 


cribriform plate 
small, flattened areas with numerous small openings, located to either 
side of the midline in the floor of the anterior cranial fossa; formed by 
the ethmoid bone 


crista galli 
small upward projection located at the midline in the floor of the 
anterior cranial fossa; formed by the ethmoid bone 


ethmoid air cell 
one of several small, air-filled spaces located within the lateral sides of 
the ethmoid bone, between the orbit and upper nasal cavity 


ethmoid bone 
unpaired bone that forms the roof and upper, lateral walls of the nasal 
cavity, portions of the floor of the anterior cranial fossa and medial 
wall of orbit, and the upper portion of the nasal septum 


external acoustic meatus 
ear canal opening located on the lateral side of the skull 


external occipital protuberance 
small bump located at the midline on the posterior skull 


facial bones 


fourteen bones that support the facial structures and form the upper 
and lower jaws and the hard palate 


foramen lacerum 
irregular opening in the base of the skull, located inferior to the exit of 
carotid canal 


foramen magnum 
large opening in the occipital bone of the skull through which the 
spinal cord emerges and the vertebral arteries enter the cranium 


foramen ovale of the middle cranial fossa 
oval-shaped opening in the floor of the middle cranial fossa 


foramen rotundum 
round opening in the floor of the middle cranial fossa, located between 
the superior orbital fissure and foramen ovale 


foramen spinosum 
small opening in the floor of the middle cranial fossa, located lateral to 
the foramen ovale 


frontal bone 
unpaired bone that forms forehead, roof of orbit, and floor of anterior 
cranial fossa 


frontal sinus 
air-filled space within the frontal bone; most anterior of the paranasal 
sinuses 


glabella 
slight depression of frontal bone, located at the midline between the 
eyebrows 


greater wings of sphenoid bone 
lateral projections of the sphenoid bone that form the anterior wall of 
the middle cranial fossa and an area of the lateral skull 


hard palate 
bony structure that forms the roof of the mouth and floor of the nasal 
cavity, formed by the palatine process of the maxillary bones and the 
horizontal plate of the palatine bones 


horizontal plate 
medial extension from the palatine bone that forms the posterior 
quarter of the hard palate 


hypoglossal canal 
paired openings that pass anteriorly from the anterior-lateral margins 
of the foramen magnum deep to the occipital condyles 


hypophyseal (pituitary) fossa 
shallow depression on top of the sella turcica that houses the pituitary 
(hypophyseal) gland 


inferior nasal concha 
one of the paired bones that project from the lateral walls of the nasal 
cavity to form the largest and most inferior of the nasal conchae 


infraorbital foramen 
opening located on anterior skull, below the orbit 


infratemporal fossa 
space on lateral side of skull, below the level of the zygomatic arch 
and deep (medial) to the ramus of the mandible 


internal acoustic meatus 
opening into petrous ridge, located on the lateral wall of the posterior 
cranial fossa 


jugular foramen 
irregularly shaped opening located in the lateral floor of the posterior 
cranial cavity 


lacrimal bone 
paired bones that contribute to the anterior-medial wall of each orbit 


lacrimal fossa 
shallow depression in the anterior-medial wall of the orbit, formed by 
the lacrimal bone that gives rise to the nasolacrimal canal 


lambdoid suture 
inverted V-shaped joint that unites the occipital bone to the right and 
left parietal bones on the posterior skull 


lateral pterygoid plate 
paired, flattened bony projections of the sphenoid bone located on the 
inferior skull, lateral to the medial pterygoid plate 


lesser wings of the sphenoid bone 
lateral extensions of the sphenoid bone that form the bony lip 
separating the anterior and middle cranial fossae 


lingula 
small flap of bone located on the inner (medial) surface of mandibular 
ramus, next to the mandibular foramen 


mandible 
unpaired bone that forms the lower jaw bone; the only moveable bone 
of the skull 


mandibular foramen 
opening located on the inner (medial) surface of the mandibular ramus 


mandibular fossa 
oval depression located on the inferior surface of the skull 


mandibular notch 
large U-shaped notch located between the condylar process and 
coronoid process of the mandible 


mastoid process 
large bony prominence on the inferior, lateral skull, just behind the 
earlobe 


maxillary bone 
(also, maxilla) paired bones that form the upper jaw and anterior 
portion of the hard palate 


maxillary sinus 
air-filled space located with each maxillary bone; largest of the 
paranasal sinuses 


medial pterygoid plate 
paired, flattened bony projections of the sphenoid bone located on the 
inferior skull medial to the lateral pterygoid plate; form the posterior 
portion of the nasal cavity lateral wall 


mental foramen 
opening located on the anterior-lateral side of the mandibular body 


mental protuberance 
inferior margin of anterior mandible that forms the chin 


middle cranial fossa 
centrally located cranial fossa that extends from the lesser wings of the 
sphenoid bone to the petrous ridge 


middle nasal concha 
nasal concha formed by the ethmoid bone that is located between the 
superior and inferior conchae 


mylohyoid line 
bony ridge located along the inner (medial) surface of the mandibular 
body 


nasal bone 
paired bones that form the base of the nose 


nasal cavity 
opening through skull for passage of air 


nasal conchae 


curved bony plates that project from the lateral walls of the nasal 
cavity; include the superior and middle nasal conchae, which are parts 
of the ethmoid bone, and the independent inferior nasal conchae bone 


nasal septum 
flat, midline structure that divides the nasal cavity into halves, formed 
by the perpendicular plate of the ethmoid bone, vomer bone, and septal 
cartilage 


nasolacrimal canal 
passage for drainage of tears that extends downward from the medial- 
anterior orbit to the nasal cavity, terminating behind the inferior nasal 
conchae 


occipital bone 
unpaired bone that forms the posterior portions of the brain case and 
base of the skull 


occipital condyle 
paired, oval-shaped bony knobs located on the inferior skull, to either 
side of the foramen magnum 


optic canal 
opening spanning between middle cranial fossa and posterior orbit 


orbit 
bony socket that contains the eyeball and associated muscles 


palatine bone 
paired bones that form the posterior quarter of the hard palate and a 
small area in floor of the orbit 


palatine process 
medial projection from the maxilla bone that forms the anterior three 
quarters of the hard palate 


paranasal sinuses 


cavities within the skull that are connected to the conchae that serve to 


warm and humidify incoming air, produce mucus, and lighten the 
weight of the skull; consist of frontal, maxillary, sphenoidal, and 
ethmoidal sinuses 


parietal bone 
paired bones that form the upper, lateral sides of the skull 


perpendicular plate of the ethmoid bone 
downward, midline extension of the ethmoid bone that forms the 
superior portion of the nasal septum 


petrous ridge 
petrous portion of the temporal bone that forms a large, triangular 
ridge in the floor of the cranial cavity, separating the middle and 
posterior cranial fossae; houses the middle and inner ear structures 


posterior cranial fossa 
deepest and most posterior cranial fossa; extends from the petrous 
ridge to the occipital bone 


pterion 
H-shaped suture junction region that unites the frontal, parietal, 
temporal, and sphenoid bones on the lateral side of the skull 


ramus of the mandible 
vertical portion of the mandible 


Sagittal suture 
joint that unites the right and left parietal bones at the midline along 
the top of the skull 


sella turcica 


elevated area of sphenoid bone located at midline of the middle cranial 


fossa 


septal cartilage 


flat cartilage structure that forms the anterior portion of the nasal 
septum 


sphenoid bone 
unpaired bone that forms the central base of skull 


sphenoid sinus 
air-filled space located within the sphenoid bone; most posterior of the 
paranasal sinuses 


squamous suture 
joint that unites the parietal bone to the squamous portion of the 
temporal bone on the lateral side of the skull 


styloid process 
downward projecting, elongated bony process located on the inferior 
aspect of the skull 


stylomastoid foramen 
opening located on inferior skull, between the styloid process and 
mastoid process 


superior nasal concha 
smallest and most superiorly located of the nasal conchae; formed by 
the ethmoid bone 


superior nuchal line 
paired bony lines on the posterior skull that extend laterally from the 
external occipital protuberance 


superior orbital fissure 
irregularly shaped opening between the middle cranial fossa and the 
posterior orbit 


supraorbital foramen 
opening located on anterior skull, at the superior margin of the orbit 


supraorbital margin 


superior margin of the orbit 


suture 
junction line at which adjacent bones of the skull are united by fibrous 
connective tissue 


temporal bone 
paired bones that form the lateral, inferior portions of the skull, with 
Squamous, mastoid, and petrous portions 


temporal fossa 
shallow space on the lateral side of the skull, above the level of the 
zygomatic arch 


temporal process of the zygomatic bone 
short extension from the zygomatic bone that forms the anterior 
portion of the zygomatic arch 


vomer bone 
unpaired bone that forms the inferior and posterior portions of the 
nasal septum 


zygomatic arch 
elongated, free-standing arch on the lateral skull, formed anteriorly by 
the temporal process of the zygomatic bone and posteriorly by the 
zygomatic process of the temporal bone 


zygomatic bone 
cheekbone; paired bones that contribute to the lateral orbit and anterior 
zygomatic arch 


zygomatic process of the temporal bone 
extension from the temporal bone that forms the posterior portion of 
the zygomatic arch 


The Vertebral Column 
By the end of this section, you will be able to: 


Describe each region of the vertebral column and the number of bones 
in each region 

Discuss the curves of the vertebral column and how these change after 
birth 

Describe a typical vertebra and determine the distinguishing 
characteristics for vertebrae in each vertebral region and features of the 
sacrum and the coccyx 

Define the structure of an intervertebral disc 

Determine the location of the ligaments that provide support for the 
vertebral column 


The vertebral column is also known as the spinal column or spine ([link]). It 
consists of a sequence of vertebrae (singular = vertebra), each of which is 
separated and united by an intervertebral disc. Together, the vertebrae and 
intervertebral discs form the vertebral column. It is a flexible column that 
supports the head, neck, and body and allows for their movements. It also 
protects the spinal cord, which passes down the back through openings in 
the vertebrae. 

Vertebral Column 


7 Cervical vertebrae 
(C1-C7) form cervical curve 


12 Thoracic vertbrae 
(T1-T12) form thoracic curve 


5 Lumbar vertebrae (L1—L5) 
form lumbar curve 


Fused vertebrae of sacrum 
and coccyx form 
sacrococcygeal curve 


The adult vertebral column consists of 24 vertebrae, 
plus the sacrum and coccyx. The vertebrae are divided 
into three regions: cervical C1—C7 vertebrae, thoracic 
T1—-T12 vertebrae, and lumbar L1—L5 vertebrae. The 

vertebral column is curved, with two primary 
curvatures (thoracic and sacrococcygeal curves) and 
two secondary curvatures (cervical and lumbar 
curves). 


Regions of the Vertebral Column 


The vertebral column originally develops as a series of 33 vertebrae, but 
this number is eventually reduced to 24 vertebrae, plus the sacrum and 
coccyx. The vertebral column is subdivided into five regions, with the 


vertebrae in each area named for that region and numbered in descending 
order. In the neck, there are seven cervical vertebrae, each designated with 
the letter “C” followed by its number. Superiorly, the C1 vertebra articulates 
(forms a joint) with the occipital condyles of the skull. Inferiorly, C1 
articulates with the C2 vertebra, and so on. Below these are the 12 thoracic 
vertebrae, designated T1—T12. The lower back contains the L1—L5 lumbar 
vertebrae. The single sacrum, which is also part of the pelvis, is formed by 
the fusion of five sacral vertebrae. Similarly, the coccyx, or tailbone, results 
from the fusion of four small coccygeal vertebrae. However, the sacral and 
coccygeal fusions do not start until age 20 and are not completed until 
middle age. 


An interesting anatomical fact is that almost all mammals have seven 
cervical vertebrae, regardless of body size. This means that there are large 
variations in the size of cervical vertebrae, ranging from the very small 
cervical vertebrae of a shrew to the greatly elongated vertebrae in the neck 
of a giraffe. In a full-grown giraffe, each cervical vertebra is 11 inches tall. 


Curvatures of the Vertebral Column 


The adult vertebral column does not form a straight line, but instead has 
four curvatures along its length (see [link]). These curves increase the 
vertebral column’s strength, flexibility, and ability to absorb shock. When 
the load on the spine is increased, by carrying a heavy backpack for 
example, the curvatures increase in depth (become more curved) to 
accommodate the extra weight. They then spring back when the weight is 
removed. The four adult curvatures are classified as either primary or 
secondary curvatures. Primary curves are retained from the original fetal 
curvature, while secondary curvatures develop after birth. 


During fetal development, the body is flexed anteriorly into the fetal 
position, giving the entire vertebral column a single curvature that is 
concave anteriorly. In the adult, this fetal curvature is retained in two 
regions of the vertebral column as the thoracic curve, which involves the 
thoracic vertebrae, and the sacrococcygeal curve, formed by the sacrum 
and coccyx. Each of these is thus called a primary curve because they are 
retained from the original fetal curvature of the vertebral column. 


A secondary curve develops gradually after birth as the child learns to sit 
upright, stand, and walk. Secondary curves are concave posteriorly, 
opposite in direction to the original fetal curvature. The cervical curve of 
the neck region develops as the infant begins to hold their head upright 
when sitting. Later, as the child begins to stand and then to walk, the 
lumbar curve of the lower back develops. In adults, the lumbar curve is 
generally deeper in females. 


Disorders associated with the curvature of the spine include kyphosis (an 
excessive posterior curvature of the thoracic region), lordosis (an excessive 
anterior curvature of the lumbar region), and scoliosis (an abnormal, lateral 
curvature, accompanied by twisting of the vertebral column). 


Note: 

Disorders of the... 

Vertebral Column 

Developmental anomalies, pathological changes, or obesity can enhance 
the normal vertebral column curves, resulting in the development of 
abnormal or excessive curvatures ({link]). Kyphosis, also referred to as 
humpback or hunchback, is an excessive posterior curvature of the thoracic 
region. This can develop when osteoporosis causes weakening and erosion 
of the anterior portions of the upper thoracic vertebrae, resulting in their 
gradual collapse ({link]). Lordosis, or swayback, is an excessive anterior 
curvature of the lumbar region and is most commonly associated with 
obesity or late pregnancy. The accumulation of body weight in the 
abdominal region results an anterior shift in the line of gravity that carries 
the weight of the body. This causes in an anterior tilt of the pelvis and a 
pronounced enhancement of the lumbar curve. 

Scoliosis is an abnormal, lateral curvature, accompanied by twisting of the 
vertebral column. Compensatory curves may also develop in other areas of 
the vertebral column to help maintain the head positioned over the feet. 
Scoliosis is the most common vertebral abnormality among girls. The 
cause is usually unknown, but it may result from weakness of the back 
muscles, defects such as differential growth rates in the right and left sides 
of the vertebral column, or differences in the length of the lower limbs. 


When present, scoliosis tends to get worse during adolescent growth 
spurts. Although most individuals do not require treatment, a back brace 
may be recommended for growing children. In extreme cases, surgery may 
be required. 

Excessive vertebral curves can be identified while an individual stands in 
the anatomical position. Observe the vertebral profile from the side and 
then from behind to check for kyphosis or lordosis. Then have the person 
bend forward. If scoliosis is present, an individual will have difficulty in 
bending directly forward, and the right and left sides of the back will not 
be level with each other in the bent position. 

Abnormal Curvatures of the Vertebral Column 


(a) Scoliosis (b) Kyphosis (c) Lordosis 


(a) Scoliosis is an abnormal lateral bending of the vertebral 
column. (b) An excessive curvature of the upper thoracic 
vertebral column is called kyphosis. (c) Lordosis is an 
excessive curvature in the lumbar region of the vertebral 
column. 


Osteoporosis 


Normal Bone loss 
vertebrae amplifies curvature 


Osteoporosis is an age-related 
disorder that causes the gradual 
loss of bone density and strength. 
When the thoracic vertebrae are 
affected, there can be a gradual 
collapse of the vertebrae. This 
results in kyphosis, an excessive 
curvature of the thoracic region. 


Note: 


— 
meee OPENStAX COLLEGE 


Osteoporosis is a common age-related bone disease in which bone density 
and strength is decreased. Watch this video to get a better understanding of 
how thoracic vertebrae may become weakened and may fracture due to this 
disease. How may vertebral osteoporosis contribute to kyphosis? 


General Structure of a Vertebra 


Within the different regions of the vertebral column, vertebrae vary in size 
and shape, but they all follow a similar structural pattern. A typical vertebra 
will consist of a body, a vertebral arch, and seven processes ({link]). 


The body is the anterior portion of each vertebra and is the part that 
supports the body weight. Because of this, the vertebral bodies 
progressively increase in size and thickness going down the vertebral 
column. The bodies of adjacent vertebrae are separated and strongly united 
by an intervertebral disc. 


The vertebral arch forms the posterior portion of each vertebra. It consists 
of four parts, the right and left pedicles and the right and left laminae. Each 
pedicle forms one of the lateral sides of the vertebral arch. The pedicles are 
anchored to the posterior side of the vertebral body. Each lamina forms part 
of the posterior roof of the vertebral arch. The large opening between the 
vertebral arch and body is the vertebral foramen, which contains the 
spinal cord. In the intact vertebral column, the vertebral foramina of all of 
the vertebrae align to form the vertebral (spinal) canal, which serves as 
the bony protection and passageway for the spinal cord down the back. 
When the vertebrae are aligned together in the vertebral column, notches in 
the margins of the pedicles of adjacent vertebrae together form an 
intervertebral foramen, the opening through which a spinal nerve exits 
from the vertebral column ([link]). 


Seven processes arise from the vertebral arch. Each paired transverse 
process projects laterally and arises from the junction point between the 
pedicle and lamina. The single spinous process (vertebral spine) projects 
posteriorly at the midline of the back. The vertebral spines can easily be felt 
as a Series of bumps just under the skin down the middle of the back. The 
transverse and spinous processes serve as important muscle attachment 
sites. A superior articular process extends or faces upward, and an 
inferior articular process faces or projects downward on each side of a 
vertebrae. The paired superior articular processes of one vertebra join with 
the corresponding paired inferior articular processes from the next higher 
vertebra. These junctions form slightly moveable joints between the 


adjacent vertebrae. The shape and orientation of the articular processes vary 
in different regions of the vertebral column and play a major role in 
determining the type and range of motion available in each region. 

Parts of a Typical Vertebra 


Anterior Posterior 
Posterior 


Spinal cord 


Spinal cord 
Spinous 
process 


Transverse 
process 


Facet of superior 
articular process 


Vertebral 
foramen 


Intervetebral disc 


Facet for head 


Vertebral arch: of rib 


Lamina 


Facet of 
superior 
articular 
process 


Inferior articular 
process 


Facet for 


head of rib Spinal nerve exiting 


Anterior through intervertebral 


foramen Spinous process 


Superior view Left posterolateral view 
of articulated vertebrae 


A typical vertebra consists of a body and a vertebral arch. The 
arch is formed by the paired pedicles and paired laminae. 
Arising from the vertebral arch are the transverse, spinous, 
superior articular, and inferior articular processes. The 
vertebral foramen provides for passage of the spinal cord. Each 
spinal nerve exits through an intervertebral foramen, located 
between adjacent vertebrae. Intervertebral discs unite the 
bodies of adjacent vertebrae. 


Intervertebral Disc 


Vertebral body 


Intervertebral foramen 


o> Anulus fibrosus ~~ 


Nucleus pulposus 


Lateral view Superior view 


The bodies of adjacent vertebrae are separated and 
united by an intervertebral disc, which provides 
padding and allows for movements between adjacent 
vertebrae. The disc consists of a fibrous outer layer 
called the anulus fibrosus and a gel-like center called 
the nucleus pulposus. The intervertebral foramen is the 
opening formed between adjacent vertebrae for the 
exit of a spinal nerve. 


Regional Modifications of Vertebrae 


In addition to the general characteristics of a typical vertebra described 
above, vertebrae also display characteristic size and structural features that 
vary between the different vertebral column regions. Thus, cervical 
vertebrae are smaller than lumbar vertebrae due to differences in the 
proportion of body weight that each supports. Thoracic vertebrae have sites 
for rib attachment, and the vertebrae that give rise to the sacrum and coccyx 
have fused together into single bones. 


Cervical Vertebrae 


Typical cervical vertebrae, such as C4 or C5, have several characteristic 
features that differentiate them from thoracic or lumbar vertebrae ([link]). 
Cervical vertebrae have a small body, reflecting the fact that they carry the 
least amount of body weight. Cervical vertebrae usually have a bifid (Y- 
shaped) spinous process. The spinous processes of the C3—C6 vertebrae are 
short, but the spine of C7 is much longer. You can find these vertebrae by 
running your finger down the midline of the posterior neck until you 
encounter the prominent C7 spine located at the base of the neck. The 
transverse processes of the cervical vertebrae are sharply curved (U-shaped) 
to allow for passage of the cervical spinal nerves. Each transverse process 
also has an opening called the transverse foramen. An important artery 
that supplies the brain ascends up the neck by passing through these 


openings. The superior and inferior articular processes of the cervical 
vertebrae are flattened and largely face upward or downward, respectively. 


The first and second cervical vertebrae are further modified, giving each a 
distinctive appearance. The first cervical (C1) vertebra is also called the 
atlas, because this is the vertebra that supports the skull on top of the 
vertebral column (in Greek mythology, Atlas was the god who supported 
the heavens on his shoulders). The C1 vertebra does not have a body or 
spinous process. Instead, it is ring-shaped, consisting of an anterior arch 
and a posterior arch. The transverse processes of the atlas are longer and 
extend more laterally than do the transverse processes of any other cervical 
vertebrae. The superior articular processes face upward and are deeply 
curved for articulation with the occipital condyles on the base of the skull. 
The inferior articular processes are flat and face downward to join with the 
superior articular processes of the C2 vertebra. 


The second cervical (C2) vertebra is called the axis, because it serves as the 
axis for rotation when turning the head toward the right or left. The axis 
resembles typical cervical vertebrae in most respects, but is easily 
distinguished by the dens (odontoid process), a bony projection that extends 
upward from the vertebral body. The dens joins with the inner aspect of the 
anterior arch of the atlas, where it is held in place by transverse ligament. 
Cervical Vertebrae 


Dens of axis 
Transverse 


ligament s 
Spinous process (bifid) & C4 f 
SS ers 
Vertebral foramen C; (atlas) “> ' 


Cz (axis) —— > 
Pedicle C3 : 


Lamina Inferior 


articular 
process 


Superior 
articular 
process Bifid spinous 


process 


ieee? Transverse 
process 
process 
Transverse C; (vertebra 
foramen Body Groove prominens) 
for spinal 
nerve 


Structure of a typical cervical vertebra 


Dens 
Superior articular Transverse 
facet fie process 
Dens - 
Superior articular Anterior arch Transverse 
foramen 
facet 


Transverse 


Lamina 
process 


Transverse 
foramen 


Posterior arch Spinous process 


Ligament 
Superior view of atlas Superior view of axis 


Dens 


Transverse 
process 


Inferior articular Body 
process 


Anterior view of axis 


A typical cervical vertebra has a small body, a bifid spinous 
process, transverse processes that have a transverse foramen 
and are curved for spinal nerve passage. The atlas (C1 
vertebra) does not have a body or spinous process. It consists 
of an anterior and a posterior arch and elongated transverse 
processes. The axis (C2 vertebra) has the upward projecting 
dens, which articulates with the anterior arch of the atlas. 


Thoracic Vertebrae 


The bodies of the thoracic vertebrae are larger than those of cervical 
vertebrae ({link]). The characteristic feature for a typical midthoracic 
vertebra is the spinous process, which is long and has a pronounced 
downward angle that causes it to overlap the next inferior vertebra. The 
superior articular processes of thoracic vertebrae face anteriorly and the 
inferior processes face posteriorly. These orientations are important 
determinants for the type and range of movements available to the thoracic 
region of the vertebral column. 


Thoracic vertebrae have several additional articulation sites, each of which 
is called a facet, where a rib is attached. Most thoracic vertebrae have two 
facets located on the lateral sides of the body, each of which is called a 
costal facet (costal = “rib”). These are for articulation with the head (end) 
of arib. An additional facet is located on the transverse process for 
articulation with the tubercle of a rib. 

Thoracic Vertebrae 


Superior articular 
process 


Articular facet 
for tubercle of rib 


Transverse 
process 


Pedicle 


Intervertebral 
Lamina disc 
Body 


Spinous 


rocess ' 
P Superior costal 


facet 


fe 
vy] // \nferior Inferior costal 
iff articular facet 


process 


A typical thoracic vertebra is distinguished by the 


spinous process, which is long and projects 
downward to overlap the next inferior vertebra. It 
also has articulation sites (facets) on the vertebral 
body and a transverse process for rib attachment. 


Rib Articulation in Thoracic Vertebrae 
Superior L 
articular facets U a 


Superior costal facet 


7 


fi) 
Crea ZY 
\ Z - 
Facet for —_ WAN eS 
; \\ YYY 
tubercle of rib 


Ay Body of vertebra 


| 
we tl Head of rib 
Zl >: b 


Ns Intervertebral disc 
Tranverse 


r ; 
processes Fit— Neck of rib 
él 


{ ! 
Se Body of vertebra 
; 7 


= Tubercle of rib 
BS . 
. Inferior costal facet 


Spinous 
process 


Thoracic vertebrae have superior and 
inferior articular facets on the vertebral 
body for articulation with the head of a 

rib, and a transverse process facet for 

articulation with the rib tubercle. 


Lumbar Vertebrae 


Lumbar vertebrae carry the greatest amount of body weight and are thus 
characterized by the large size and thickness of the vertebral body ((link]). 
They have short transverse processes and a short, blunt spinous process that 
projects posteriorly. The articular processes are large, with the superior 
process facing backward and the inferior facing forward. 


Lumbar Vertebrae 


Superior articular 


Transverse 
process 


Inferior 


articular 
process 

; go ST, 
Spinous ————4—“ 

: [7 


process 


Intervertebral 
disc 


Inferior articular process 


Lumbar vertebrae are characterized by 
having a large, thick body and a short, 
rounded spinous process. 


Sacrum and Coccyx 


The sacrum is a triangular-shaped bone that is thick and wide across its 
superior base where it is weight bearing and then tapers down to an inferior, 
non-weight bearing apex ([link]). It is formed by the fusion of five sacral 
vertebrae, a process that does not begin until after the age of 20. On the 
anterior surface of the older adult sacrum, the lines of vertebral fusion can 
be seen as four transverse ridges. On the posterior surface, running down 
the midline, is the median sacral crest, a bumpy ridge that is the remnant 
of the fused spinous processes (median = “midline”; while medial = 
“toward, but not necessarily at, the midline”). Similarly, the fused 
transverse processes of the sacral vertebrae form the lateral sacral crest. 


The sacral promontory is the anterior lip of the superior base of the 
sacrum. Lateral to this is the roughened auricular surface, which joins with 
the ilium portion of the hipbone to form the immobile sacroiliac joints of 
the pelvis. Passing inferiorly through the sacrum is a bony tunnel called the 
sacral canal, which terminates at the sacral hiatus near the inferior tip of 
the sacrum. The anterior and posterior surfaces of the sacrum have a series 
of paired openings called sacral foramina (singular = foramen) that 
connect to the sacral canal. Each of these openings is called a posterior 
(dorsal) sacral foramen or anterior (ventral) sacral foramen. These 
openings allow for the anterior and posterior branches of the sacral spinal 
nerves to exit the sacrum. The superior articular process of the sacrum, 
one of which is found on either side of the superior opening of the sacral 
canal, articulates with the inferior articular processes from the L5 vertebra. 


The coccyx, or tailbone, is derived from the fusion of four very small 
coccygeal vertebrae (see [link]). It articulates with the inferior tip of the 
sacrum. It is not weight bearing in the standing position, but may receive 
some body weight when sitting. 

Sacrum and Coccyx 


Sacral Body Facet of superior 
Sacral promontory 


articular process 


Body of first 
sacral vertebra wi 
T 


Transverse ridges 
(sites of vertebral 
fusion) 


Auricular 
surface 


Lateral 
sacral 
crest 


Posterior 
sacral 
foramina 


Anterior sacral 


foramina A Sacral hiatus 


- Coccyx 


Anterior view Posterior view 


Apex 


Coccyx 


The sacrum is formed from the fusion of five sacral vertebrae, 
whose lines of fusion are indicated by the transverse ridges. 
The fused spinous processes form the median sacral crest, 
while the lateral sacral crest arises from the fused transverse 
processes. The coccyx is formed by the fusion of four small 
coccygeal vertebrae. 


Intervertebral Discs and Ligaments of the Vertebral Column 


The bodies of adjacent vertebrae are strongly anchored to each other by an 
intervertebral disc. This structure provides padding between the bones 
during weight bearing, and because it can change shape, also allows for 
movement between the vertebrae. Although the total amount of movement 
available between any two adjacent vertebrae is small, when these 
movements are summed together along the entire length of the vertebral 
column, large body movements can be produced. Ligaments that extend 
along the length of the vertebral column also contribute to its overall 
support and stability. 


Intervertebral Disc 


An intervertebral disc is a fibrocartilaginous pad that fills the gap between 
adjacent vertebral bodies (see [link]). Each disc is anchored to the bodies of 
its adjacent vertebrae, thus strongly uniting these. The discs also provide 
padding between vertebrae during weight bearing. Because of this, 
intervertebral discs are thin in the cervical region and thickest in the lumbar 
region, which carries the most body weight. In total, the intervertebral discs 
account for approximately 25 percent of your body height between the top 
of the pelvis and the base of the skull. Intervertebral discs are also flexible 
and can change shape to allow for movements of the vertebral column. 


Each intervertebral disc consists of two parts. The anulus fibrosus is the 
tough, fibrous outer layer of the disc. It forms a circle (anulus = “ring” or 
“circle”) and is firmly anchored to the outer margins of the adjacent 
vertebral bodies. Inside is the nucleus pulposus, consisting of a softer, 
more gel-like material. It has a high water content that serves to resist 
compression and thus is important for weight bearing. With increasing age, 
the water content of the nucleus pulposus gradually declines. This causes 
the disc to become thinner, decreasing total body height somewhat, and 


reduces the flexibility and range of motion of the disc, making bending 
more difficult. 


The gel-like nature of the nucleus pulposus also allows the intervertebral 
disc to change shape as one vertebra rocks side to side or forward and back 
in relation to its neighbors during movements of the vertebral column. 
Thus, bending forward causes compression of the anterior portion of the 
disc but expansion of the posterior disc. If the posterior anulus fibrosus is 
weakened due to injury or increasing age, the pressure exerted on the disc 
when bending forward and lifting a heavy object can cause the nucleus 
pulposus to protrude posteriorly through the anulus fibrosus, resulting in a 
herniated disc (“ruptured” or “slipped” disc) ({link]). The posterior bulging 
of the nucleus pulposus can cause compression of a spinal nerve at the point 
where it exits through the intervertebral foramen, with resulting pain and/or 
muscle weakness in those body regions supplied by that nerve. The most 
common sites for disc herniation are the L4/L5 or L5/S1 intervertebral 
discs, which can cause sciatica, a widespread pain that radiates from the 
lower back down the thigh and into the leg. Similar injuries of the C5/C6 or 
C6/C7 intervertebral discs, following forcible hyperflexion of the neck from 
a collision accident or football injury, can produce pain in the neck, 
shoulder, and upper limb. 

Herniated Intervertebral Disc 


Spinal cord within 
vertebral canal 


Herniated disc 
compresses 
nerve in 
intervertebral 


foramen 
Nucleus 


pulposus \ - 


Anulus 
fibrosus 


Herniated 
portion of disc 


Superior view 


Weakening of the anulus fibrosus can result in 
herniation (protrusion) of the nucleus pulposus and 
compression of a spinal nerve, resulting in pain 


and/or muscle weakness in the body regions 
supplied by that nerve. 


Note: 
os 
1 ‘all 
at t 
4 openstax COLLEGE 
er 
[my: i C 


Watch this animation to see what it means to “slip” a disk. Watch this 
second animation to see one possible treatment for a herniated disc, 
removing and replacing the damaged disc with an artificial one that allows 
for movement between the adjacent certebrae. How could lifting a heavy 
object produce pain in a lower limb? 


Ligaments of the Vertebral Column 


Adjacent vertebrae are united by ligaments that run the length of the 
vertebral column along both its posterior and anterior aspects ([link]). These 
serve to resist excess forward or backward bending movements of the 
vertebral column, respectively. 


The anterior longitudinal ligament runs down the anterior side of the 
entire vertebral column, uniting the vertebral bodies. It serves to resist 
excess backward bending of the vertebral column. Protection against this 
movement is particularly important in the neck, where extreme posterior 
bending of the head and neck can stretch or tear this ligament, resulting in a 
painful whiplash injury. Prior to the mandatory installation of seat 
headrests, whiplash injuries were common for passengers involved in a 
rear-end automobile collision. 


The supraspinous ligament is located on the posterior side of the vertebral 
column, where it interconnects the spinous processes of the thoracic and 
lumbar vertebrae. This strong ligament supports the vertebral column 
during forward bending motions. In the posterior neck, where the cervical 
spinous processes are short, the supraspinous ligament expands to become 
the nuchal ligament (nuchae = “nape” or “back of the neck”). The nuchal 
ligament is attached to the cervical spinous processes and extends upward 
and posteriorly to attach to the midline base of the skull, out to the external 
occipital protuberance. It supports the skull and prevents it from falling 
forward. This ligament is much larger and stronger in four-legged animals 
such as cows, where the large skull hangs off the front end of the vertebral 
column. You can easily feel this ligament by first extending your head 
backward and pressing down on the posterior midline of your neck. Then 
tilt your head forward and you will fill the nuchal ligament popping out as it 
tightens to limit anterior bending of the head and neck. 


Additional ligaments are located inside the vertebral canal, next to the 
spinal cord, along the length of the vertebral column. The posterior 
longitudinal ligament is found anterior to the spinal cord, where it is 
attached to the posterior sides of the vertebral bodies. Posterior to the spinal 
cord is the ligamentum flavum (“yellow ligament”). This consists of a 
series of short, paired ligaments, each of which interconnects the lamina 
regions of adjacent vertebrae. The ligamentum flavum has large numbers of 
elastic fibers, which have a yellowish color, allowing it to stretch and then 
pull back. Both of these ligaments provide important support for the 
vertebral column when bending forward. 

Ligaments of Vertebral Column 


External 
occipital 
protuberance 


Nuchal 
ligament 


Spinous process 
of T1 vertebra 


Anterior 
longitudinal 


Supraspinous ligament 


ligament 


The anterior longitudinal ligament 
runs the length of the vertebral 
column, uniting the anterior sides 
of the vertebral bodies. The 
supraspinous ligament connects 
the spinous processes of the 
thoracic and lumbar vertebrae. In 
the posterior neck, the 
supraspinous ligament enlarges to 
form the nuchal ligament, which 
attaches to the cervical spinous 
processes and to the base of the 
skull. 


Note: 


oe 


openstax COLLEGE 


Use this tool to identify the bones, intervertebral discs, and ligaments of 
the vertebral column. The thickest portions of the anterior longitudinal 
ligament and the supraspinous ligament are found in which regions of the 
vertebral column? 


Note: 

Career Connections 

Chiropractor 

Chiropractors are health professionals who use nonsurgical techniques to 
help patients with musculoskeletal system problems that involve the bones, 
muscles, ligaments, tendons, or nervous system. They treat problems such 
as neck pain, back pain, joint pain, or headaches. Chiropractors focus on 
the patient’s overall health and can also provide counseling related to 
lifestyle issues, such as diet, exercise, or sleep problems. If needed, they 
will refer the patient to other medical specialists. 

Chiropractors use a drug-free, hands-on approach for patient diagnosis and 
treatment. They will perform a physical exam, assess the patient’s posture 
and spine, and may perform additional diagnostic tests, including taking X- 
ray images. They primarily use manual techniques, such as spinal 
manipulation, to adjust the patient’s spine or other joints. They can 
recommend therapeutic or rehabilitative exercises, and some also include 
acupuncture, massage therapy, or ultrasound as part of the treatment 
program. In addition to those in general practice, some chiropractors 
specialize in sport injuries, neurology, orthopaedics, pediatrics, nutrition, 
internal disorders, or diagnostic imaging. 

To become a chiropractor, students must have 3—4 years of undergraduate 
education, attend an accredited, four-year Doctor of Chiropractic (D.C.) 
degree program, and pass a licensure examination to be licensed for 


practice in their state. With the aging of the baby-boom generation, 
employment for chiropractors is expected to increase. 


Chapter Review 


The vertebral column forms the neck and back. The vertebral column 
originally develops as 33 vertebrae, but is eventually reduced to 24 
vertebrae, plus the sacrum and coccyx. The vertebrae are divided into the 
cervical region (C1—C7 vertebrae), the thoracic region (T1—T12 vertebrae), 
and the lumbar region (L1—L5 vertebrae). The sacrum arises from the 
fusion of five sacral vertebrae and the coccyx from the fusion of four small 
coccygeal vertebrae. The vertebral column has four curvatures, the cervical, 
thoracic, lumbar, and sacrococcygeal curves. The thoracic and 
sacrococcygeal curves are primary curves retained from the original fetal 
curvature. The cervical and lumbar curves develop after birth and thus are 
secondary curves. The cervical curve develops as the infant begins to hold 
up the head, and the lumbar curve appears with standing and walking. 


A typical vertebra consists of an enlarged anterior portion called the body, 
which provides weight-bearing support. Attached posteriorly to the body is 
a vertebral arch, which surrounds and defines the vertebral foramen for 
passage of the spinal cord. The vertebral arch consists of the pedicles, 
which attach to the vertebral body, and the laminae, which come together to 
form the roof of the arch. Arising from the vertebral arch are the laterally 
projecting transverse processes and the posteriorly oriented spinous process. 
The superior articular processes project upward, where they articulate with 
the downward projecting inferior articular processes of the next higher 
vertebrae. 


A typical cervical vertebra has a small body, a bifid (Y-shaped) spinous 
process, and U-shaped transverse processes with a transverse foramen. In 
addition to these characteristics, the axis (C2 vertebra) also has the dens 
projecting upward from the vertebral body. The atlas (C1 vertebra) differs 
from the other cervical vertebrae in that it does not have a body, but instead 
consists of bony ring formed by the anterior and posterior arches. The atlas 
articulates with the dens from the axis. A typical thoracic vertebra is 


distinguished by its long, downward projecting spinous process. Thoracic 
vertebrae also have articulation facets on the body and transverse processes 
for attachment of the ribs. Lumbar vertebrae support the greatest amount of 
body weight and thus have a large, thick body. They also have a short, blunt 
spinous process. The sacrum is triangular in shape. The median sacral crest 
is formed by the fused vertebral spinous processes and the lateral sacral 
crest is derived from the fused transverse processes. Anterior (ventral) and 
posterior (dorsal) sacral foramina allow branches of the sacral spinal nerves 
to exit the sacrum. The auricular surfaces are articulation sites on the lateral 
sacrum that anchor the sacrum to the hipbones to form the pelvis. The 
coccyx is small and derived from the fusion of four small vertebrae. 


The intervertebral discs fill in the gaps between the bodies of adjacent 
vertebrae. They provide strong attachments and padding between the 
vertebrae. The outer, fibrous layer of a disc is called the anulus fibrosus. 
The gel-like interior is called the nucleus pulposus. The disc can change 
shape to allow for movement between vertebrae. If the anulus fibrosus is 
weakened or damaged, the nucleus pulposus can protrude outward, 
resulting in a herniated disc. 


The anterior longitudinal ligament runs along the full length of the anterior 
vertebral column, uniting the vertebral bodies. The supraspinous ligament is 
located posteriorly and interconnects the spinous processes of the thoracic 
and lumbar vertebrae. In the neck, this ligament expands to become the 
nuchal ligament. The nuchal ligament is attached to the cervical spinous 
processes and superiorly to the base of the skull, out to the external 
occipital protuberance. The posterior longitudinal ligament runs within the 
vertebral canal and unites the posterior sides of the vertebral bodies. The 
ligamentum flavum unites the lamina of adjacent vertebrae. 


Interactive Link Questions 


Exercise: 


Problem: 


Osteoporosis is a common age-related bone disease in which bone 
density and strength is decreased. Watch this video to get a better 
understanding of how thoracic vertebrae may become weakened and 
may fractured due to this disease. How may vertebral osteoporosis 
contribute to kyphosis? 


Solution: 


Osteoporosis causes thinning and weakening of the vertebral bodies. 
When this occurs in thoracic vertebrae, the bodies may collapse 
producing kyphosis, an enhanced anterior curvature of the thoracic 
vertebral column. 


Exercise: 


Problem: 


Watch this animation to see what it means to “slip” a disk. Watch this 
second animation to see one possible treatment for a herniated disc, 
removing and replacing the damaged disc with an artificial one that 
allows for movement between the adjacent certebrae. How could 
lifting a heavy object produce pain in a lower limb? 


Solution: 


Lifting a heavy object can cause an intervertebral disc in the lower 
back to bulge and compress a spinal nerve as it exits through the 
intervertebral foramen, thus producing pain in those regions of the 
lower limb supplied by that nerve. 


Exercise: 
Problem: 
Use this tool to identify the bones, intervertebral discs, and ligaments 
of the vertebral column. The thickest portions of the anterior 


longitudinal ligament and the supraspinous ligament are found in 
which regions of the vertebral column? 


Solution: 


The anterior longitudinal ligament is thickest in the thoracic region of 
the vertebral column, while the supraspinous ligament is thickest in the 
lumbar region. 


Review Questions 


Exercise: 


Problem: 
The cervical region of the vertebral column consists of 


a. seven vertebrae 

b. 12 vertebrae 

c. five vertebrae 

d. a single bone derived from the fusion of five vertebrae 


Solution: 
A 


Exercise: 


Problem:The primary curvatures of the vertebral column 


a. include the lumbar curve 

b. are remnants of the original fetal curvature 
c. include the cervical curve 

d. develop after the time of birth 


Solution: 


B 


Exercise: 


Problem: A typical vertebra has 


a. a vertebral foramen that passes through the body 

b. a superior articular process that projects downward to articulate 
with the superior portion of the next lower vertebra 

c. lamina that spans between the transverse process and spinous 
process 

d. a pair of laterally projecting spinous processes 


Solution: 


C 


Exercise: 


Problem:A typical lumbar vertebra has 


a. a Short, rounded spinous process 
b. a bifid spinous process 

c. articulation sites for ribs 

d. a transverse foramen 


Solution: 


A 
Exercise: 


Problem: 
Which is found only in the cervical region of the vertebral column? 


a. nuchal ligament 
b. ligamentum flavum 
c. supraspinous ligament 


d. anterior longitudinal ligament 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: Describe the vertebral column and define each region. 
Solution: 


Answer: The adult vertebral column consists of 24 vertebrae, plus the 
sacrum and coccyx. The vertebrae are subdivided into cervical, 
thoracic, and lumbar regions. There are seven cervical vertebrae (C1— 
C7), 12 thoracic vertebrae (T1—T12), and five lumbar vertebrae (L1— 
L5). The sacrum is derived from the fusion of five sacral vertebrae and 
the coccyx is formed by the fusion of four small coccygeal vertebrae. 


Exercise: 


Problem: Describe a typical vertebra. 
Solution: 


A typical vertebra consists of an anterior body and a posterior vertebral 
arch. The body serves for weight bearing. The vertebral arch surrounds 
and protects the spinal cord. The vertebral arch is formed by the 
pedicles, which are attached to the posterior side of the vertebral body, 
and the lamina, which come together to form the top of the arch. A 
pair of transverse processes extends laterally from the vertebral arch, at 
the junction between each pedicle and lamina. The spinous process 
extends posteriorly from the top of the arch. A pair of superior 
articular processes project upward and a pair of inferior articular 


processes project downward. Together, the notches found in the 
margins of the pedicles of adjacent vertebrae form an intervertebral 
foramen. 


Exercise: 


Problem: Describe the sacrum. 


Solution: 


The sacrum is a single, triangular-shaped bone formed by the fusion of 
five sacral vertebrae. On the posterior sacrum, the median sacral crest 
is derived from the fused spinous processes, and the lateral sacral crest 
results from the fused transverse processes. The sacral canal contains 
the sacral spinal nerves, which exit via the anterior (ventral) and 
posterior (dorsal) sacral foramina. The sacral promontory is the 
anterior lip. The sacrum also forms the posterior portion of the pelvis. 


Exercise: 


Problem: Describe the structure and function of an intervertebral disc. 
Solution: 


An intervertebral disc fills in the space between adjacent vertebrae, 
where it provides padding and weight-bearing ability, and allows for 
movements between the vertebrae. It consists of an outer anulus 
fibrosus and an inner nucleus pulposus. The anulus fibrosus strongly 
anchors the adjacent vertebrae to each other, and the high water 
content of the nucleus pulposus resists compression for weight bearing 
and can change shape to allow for vertebral column movements. 


Exercise: 


Problem: Define the ligaments of the vertebral column. 


Solution: 


The anterior longitudinal ligament is attached to the vertebral bodies 
on the anterior side of the vertebral column. The supraspinous 
ligament is located on the posterior side, where it interconnects the 
thoracic and lumbar spinous processes. In the posterior neck, this 
ligament expands to become the nuchal ligament, which attaches to the 
cervical spinous processes and the base of the skull. The posterior 
longitudinal ligament and ligamentum flavum are located inside the 
vertebral canal. The posterior longitudinal ligament unites the posterior 
sides of the vertebral bodies. The ligamentum flavum unites the lamina 
of adjacent vertebrae. 


Glossary 


anterior arch 
anterior portion of the ring-like C1 (atlas) vertebra 


anterior longitudinal ligament 
ligament that runs the length of the vertebral column, uniting the 
anterior aspects of the vertebral bodies 


anterior (ventral) sacral foramen 
one of the series of paired openings located on the anterior (ventral) 
side of the sacrum 


anulus fibrosus 
tough, fibrous outer portion of an intervertebral disc, which is strongly 
anchored to the bodies of the adjacent vertebrae 


atlas 
first cervical (C1) vertebra 


axis 
second cervical (C2) vertebra 


cervical curve 
posteriorly concave curvature of the cervical vertebral column region; 
a secondary curve of the vertebral column 


cervical vertebrae 
seven vertebrae numbered as C1—C7 that are located in the neck region 
of the vertebral column 


costal facet 
site on the lateral sides of a thoracic vertebra for articulation with the 
head of a rib 


dens 
bony projection (odontoid process) that extends upward from the body 
of the C2 (axis) vertebra 


facet 
small, flattened area on a bone for an articulation (joint) with another 
bone, or for muscle attachment 


inferior articular process 
bony process that extends downward from the vertebral arch of a 
vertebra that articulates with the superior articular process of the next 
lower vertebra 


intervertebral disc 
structure located between the bodies of adjacent vertebrae that strongly 
joins the vertebrae; provides padding, weight bearing ability, and 
enables vertebral column movements 


intervertebral foramen 
opening located between adjacent vertebrae for exit of a spinal nerve 


kyphosis 
(also, humpback or hunchback) excessive posterior curvature of the 
thoracic vertebral column region 


lamina 
portion of the vertebral arch on each vertebra that extends between the 


transverse and spinous process 


lateral sacral crest 


paired irregular ridges running down the lateral sides of the posterior 
sacrum that was formed by the fusion of the transverse processes from 
the five sacral vertebrae 


ligamentum flavum 
series of short ligaments that unite the lamina of adjacent vertebrae 


lordosis 
(also, swayback) excessive anterior curvature of the lumbar vertebral 
column region 


lumbar curve 
posteriorly concave curvature of the lumbar vertebral column region; a 
secondary curve of the vertebral column 


lumbar vertebrae 
five vertebrae numbered as L1—L5 that are located in lumbar region 
(lower back) of the vertebral column 


median sacral crest 
irregular ridge running down the midline of the posterior sacrum that 
was formed from the fusion of the spinous processes of the five sacral 
vertebrae 


nuchal ligament 
expanded portion of the supraspinous ligament within the posterior 
neck; interconnects the spinous processes of the cervical vertebrae and 
attaches to the base of the skull 


nucleus pulposus 
gel-like central region of an intervertebral disc; provides for padding, 
weight-bearing, and movement between adjacent vertebrae 


pedicle 
portion of the vertebral arch that extends from the vertebral body to the 


transverse process 


posterior arch 


posterior portion of the ring-like C1 (atlas) vertebra 


posterior longitudinal ligament 
ligament that runs the length of the vertebral column, uniting the 
posterior sides of the vertebral bodies 


posterior (dorsal) sacral foramen 
one of the series of paired openings located on the posterior (dorsal) 
side of the sacrum 


primary curve 
anteriorly concave curvatures of the thoracic and sacrococcygeal 
regions that are retained from the original fetal curvature of the 
vertebral column 


sacral canal 
bony tunnel that runs through the sacrum 


sacral foramina 
series of paired openings for nerve exit located on both the anterior 
(ventral) and posterior (dorsal) aspects of the sacrum 


sacral hiatus 
inferior opening and termination of the sacral canal 


sacral promontory 
anterior lip of the base (superior end) of the sacrum 


sacrococcygeal curve 
anteriorly concave curvature formed by the sacrum and coccyx; a 
primary curve of the vertebral column 


scoliosis 
abnormal lateral curvature of the vertebral column 


secondary curve 
posteriorly concave curvatures of the cervical and lumbar regions of 
the vertebral column that develop after the time of birth 


spinous process 
unpaired bony process that extends posteriorly from the vertebral arch 
of a vertebra 


superior articular process 
bony process that extends upward from the vertebral arch of a vertebra 
that articulates with the inferior articular process of the next higher 
vertebra 


superior articular process of the sacrum 
paired processes that extend upward from the sacrum to articulate 
(join) with the inferior articular processes from the L5 vertebra 


supraspinous ligament 
ligament that interconnects the spinous processes of the thoracic and 
lumbar vertebrae 


thoracic curve 
anteriorly concave curvature of the thoracic vertebral column region; a 
primary curve of the vertebral column 


thoracic vertebrae 
twelve vertebrae numbered as T1—T12 that are located in the thoracic 
region (upper back) of the vertebral column 


transverse foramen 
opening found only in the transverse processes of cervical vertebrae 


transverse process 
paired bony processes that extends laterally from the vertebral arch of 
a vertebra 


vertebral arch 
bony arch formed by the posterior portion of each vertebra that 
surrounds and protects the spinal cord 


vertebral (spinal) canal 


bony passageway within the vertebral column for the spinal cord that 
is formed by the series of individual vertebral foramina 


vertebral foramen 
opening associated with each vertebra defined by the vertebral arch 
that provides passage for the spinal cord 


The Thoracic Cage 
By the end of this section, you will be able to: 


e Discuss the components that make up the thoracic cage 
e Identify the parts of the sternum and define the sternal angle 
e Discuss the parts of a rib and rib classifications 


The thoracic cage (rib cage) forms the thorax (chest) portion of the body. It 
consists of the 12 pairs of ribs with their costal cartilages and the sternum 
({link]). The ribs are anchored posteriorly to the 12 thoracic vertebrae (T1— 
T12). The thoracic cage protects the heart and lungs. 


Thoracic Cage 
Superior Superior 
Jugular 
Clavicular a notch 
notch or —— 
Jugular notch Clavicular notch 
Manubrium Clavicle 
Sternal 4 
angle Sternum: 
Manubrium Scapula 
Body 
§ 


Sternal angle 
Body —_ 
) Xiphoid 
: process 


Costal cartilages 


Intercostal space 


Xiphoid aw 
process 
Inferior Inferior 
(a) Anterior view of sternum (b) Anterior view of skeleton of thorax 


The thoracic cage is formed by the (a) sternum and (b) 12 pairs 
of ribs with their costal cartilages. The ribs are anchored 
posteriorly to the 12 thoracic vertebrae. The sternum consists 
of the manubrium, body, and xiphoid process. The ribs are 
classified as true ribs (1—7) and false ribs (8-12). The last two 
pairs of false ribs are also known as floating ribs (11-12). 


Sternum 


The sternum is the elongated bony structure that anchors the anterior 
thoracic cage. It consists of three parts: the manubrium, body, and xiphoid 
process. The manubrium is the wider, superior portion of the sternum. The 
top of the manubrium has a shallow, U-shaped border called the jugular 
(suprasternal) notch. This can be easily felt at the anterior base of the 
neck, between the medial ends of the clavicles. The clavicular notch is the 
shallow depression located on either side at the superior-lateral margins of 
the manubrium. This is the site of the sternoclavicular joint, between the 
sternum and clavicle. The first ribs also attach to the manubrium. 


The elongated, central portion of the sternum is the body. The manubrium 
and body join together at the sternal angle, so called because the junction 
between these two components is not flat, but forms a slight bend. The 
second rib attaches to the sternum at the sternal angle. Since the first rib is 
hidden behind the clavicle, the second rib is the highest rib that can be 
identified by palpation. Thus, the sternal angle and second rib are important 
landmarks for the identification and counting of the lower ribs. Ribs 3—7 
attach to the sternal body. 


The inferior tip of the sternum is the xiphoid process. This small structure 
is cartilaginous early in life, but gradually becomes ossified starting during 
middle age. 


Ribs 


Each rib is a curved, flattened bone that contributes to the wall of the 
thorax. The ribs articulate posteriorly with the T1—-T12 thoracic vertebrae, 
and most attach anteriorly via their costal cartilages to the sternum. There 
are 12 pairs of ribs. The ribs are numbered 1—12 in accordance with the 
thoracic vertebrae. 


Parts of a Typical Rib 


The posterior end of a typical rib is called the head of the rib (see [link]). 
This region articulates primarily with the costal facet located on the body of 
the same numbered thoracic vertebra and to a lesser degree, with the costal 
facet located on the body of the next higher vertebra. Lateral to the head is 
the narrowed neck of the rib. A small bump on the posterior rib surface is 
the tubercle of the rib, which articulates with the facet located on the 
transverse process of the same numbered vertebra. The remainder of the rib 
is the body of the rib (shaft). Just lateral to the tubercle is the angle of the 
rib, the point at which the rib has its greatest degree of curvature. The 
angles of the ribs form the most posterior extent of the thoracic cage. In the 
anatomical position, the angles align with the medial border of the scapula. 
A shallow costal groove for the passage of blood vessels and a nerve is 
found along the inferior margin of each rib. 


Rib Classifications 


The bony ribs do not extend anteriorly completely around to the sternum. 
Instead, each rib ends in a costal cartilage. These cartilages are made of 
hyaline cartilage and can extend for several inches. Most ribs are then 
attached, either directly or indirectly, to the sternum via their costal 
cartilage (see [link]). The ribs are classified into three groups based on their 
relationship to the sternum. 


Ribs 1—7 are classified as true ribs (vertebrosternal ribs). The costal 
cartilage from each of these ribs attaches directly to the sternum. Ribs 8-12 
are called false ribs (vertebrochondral ribs). The costal cartilages from 
these ribs do not attach directly to the sternum. For ribs 8—10, the costal 
cartilages are attached to the cartilage of the next higher rib. Thus, the 
cartilage of rib 10 attaches to the cartilage of rib 9, rib 9 then attaches to rib 
8, and rib 8 is attached to rib 7. The last two false ribs (11-12) are also 
called floating ribs (vertebral ribs). These are short ribs that do not attach 
to the sternum at all. Instead, their small costal cartilages terminate within 
the musculature of the lateral abdominal wall. 


Chapter Review 


The thoracic cage protects the heart and lungs. It is composed of 12 pairs of 
ribs with their costal cartilages and the sternum. The ribs are anchored 
posteriorly to the 12 thoracic vertebrae. The sternum consists of the 
manubrium, body, and xiphoid process. The manubrium and body are 
joined at the sternal angle, which is also the site for attachment of the 
second ribs. 


Ribs are flattened, curved bones and are numbered 1-12. Posteriorly, the 
head of the rib articulates with the costal facets located on the bodies of 
thoracic vertebrae and the rib tubercle articulates with the facet located on 
the vertebral transverse process. The angle of the ribs forms the most 
posterior portion of the thoracic cage. The costal groove in the inferior 
margin of each rib carries blood vessels and a nerve. Anteriorly, each rib 
ends in a costal cartilage. True ribs (1—7) attach directly to the sternum via 
their costal cartilage. The false ribs (8-12) either attach to the sternum 
indirectly or not at all. Ribs 8—10 have their costal cartilages attached to the 
cartilage of the next higher rib. The floating ribs (11-12) are short and do 
not attach to the sternum or to another rib. 


Review Questions 


Exercise: 


Problem: The sternum 


a. consists of only two parts, the manubrium and xiphoid process 

b. has the sternal angle located between the manubrium and body 

c. receives direct attachments from the costal cartilages of all 12 
pairs of ribs 

d. articulates directly with the thoracic vertebrae 


Solution: 


B 


Exercise: 


Problem:The sternal angle is the 


a. junction between the body and xiphoid process 
b. site for attachment of the clavicle 

c. site for attachment of the floating ribs 

d. junction between the manubrium and body 


Solution: 


D 


Exercise: 


Problem:The tubercle of a rib 


a. is for articulation with the transverse process of a thoracic 
vertebra 

b. is for articulation with the body of a thoracic vertebra 

c. provides for passage of blood vessels and a nerve 

d. is the area of greatest rib curvature 


Solution: 


A 


Exercise: 


Problem: True ribs are 


a. ribs 8-12 

b. attached via their costal cartilage to the next higher rib 

c. made entirely of bone, and thus do not have a costal cartilage 
d. attached via their costal cartilage directly to the sternum 


Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: Define the parts and functions of the thoracic cage. 
Solution: 


The thoracic cage is formed by the 12 pairs of ribs with their costal 
cartilages and the sternum. The ribs are attached posteriorly to the 12 
thoracic vertebrae and most are anchored anteriorly either directly or 
indirectly to the sternum. The thoracic cage functions to protect the 
heart and lungs. 


Exercise: 


Problem: Describe the parts of the sternum. 
Solution: 


The sternum consists of the manubrium, body, and xiphoid process. 
The manubrium forms the expanded, superior end of the sternum. It 
has a jugular (suprasternal) notch, a pair of clavicular notches for 
articulation with the clavicles, and receives the costal cartilage of the 
first rib. The manubrium is joined to the body of the sternum at the 
sternal angle, which is also the site for attachment of the second rib 
costal cartilages. The body receives the costal cartilage attachments for 
ribs 3-7. The small xiphoid process forms the inferior tip of the 
sternum. 


Exercise: 


Problem: Discuss the parts of a typical rib. 
Solution: 


A typical rib is a flattened, curved bone. The head of a rib is attached 
posteriorly to the costal facets of the thoracic vertebrae. The rib 
tubercle articulates with the transverse process of a thoracic vertebra. 
The angle is the area of greatest rib curvature and forms the largest 
portion of the thoracic cage. The body (shaft) of a rib extends 
anteriorly and terminates at the attachment to its costal cartilage. The 
shallow costal groove runs along the inferior margin of a rib and 
carries blood vessels and a nerve. 


Exercise: 


Problem: Define the classes of ribs. 
Solution: 


Ribs are classified based on if and how their costal cartilages attach to 
the sternum. True (vertebrosternal) ribs are ribs 1—7. The costal 
cartilage for each of these attaches directly to the sternum. False 
(vertebrochondral) ribs, 8—12, are attached either indirectly or not at all 
to the sternum. Ribs 8—10 are attached indirectly to the sternum. For 
these ribs, the costal cartilage of each attaches to the cartilage of the 
next higher rib. The last false ribs (11-12) are also called floating 
(vertebral) ribs, because these ribs do not attach to the sternum at all. 
Instead, the ribs and their small costal cartilages terminate within the 
muscles of the lateral abdominal wall. 


Glossary 
angle of the rib 


portion of rib with greatest curvature; together, the rib angles form the 
most posterior extent of the thoracic cage 


body of the rib 
shaft portion of a rib 


clavicular notch 
paired notches located on the superior-lateral sides of the sternal 
manubrium, for articulation with the clavicle 


costal cartilage 
hyaline cartilage structure attached to the anterior end of each rib that 
provides for either direct or indirect attachment of most ribs to the 
sternum 


costal groove 
shallow groove along the inferior margin of a rib that provides passage 
for blood vessels and a nerve 


false ribs 
vertebrochondral ribs 8-12 whose costal cartilage either attaches 
indirectly to the sternum via the costal cartilage of the next higher rib 
or does not attach to the sternum at all 


floating ribs 
vertebral ribs 11—12 that do not attach to the sternum or to the costal 
cartilage of another rib 


head of the rib 
posterior end of a rib that articulates with the bodies of thoracic 
vertebrae 


jugular (suprasternal) notch 
shallow notch located on superior surface of sternal manubrium 


manubrium 
expanded, superior portion of the sternum 


neck of the rib 
narrowed region of a rib, next to the rib head 


sternal angle 
junction line between manubrium and body of the sternum and the site 
for attachment of the second rib to the sternum 


true ribs 
vertebrosternal ribs 1—7 that attach via their costal cartilage directly to 
the sternum 


tubercle of the rib 
small bump on the posterior side of a rib for articulation with the 
transverse process of a thoracic vertebra 


xiphoid process 
small process that forms the inferior tip of the sternum 


The Pectoral Girdle 
By the end of this section, you will be able to: 


¢ Describe the bones that form the pectoral girdle 
e List the functions of the pectoral girdle 


The appendicular skeleton includes all of the limb bones, plus the bones 
that unite each limb with the axial skeleton ({link]). The bones that attach 
each upper limb to the axial skeleton form the pectoral girdle (shoulder 
girdle). This consists of two bones, the scapula and clavicle ([link]). The 
clavicle (collarbone) is an S-shaped bone located on the anterior side of the 
shoulder. It is attached on its medial end to the sternum of the thoracic cage, 
which is part of the axial skeleton. The lateral end of the clavicle articulates 
(joins) with the scapula just above the shoulder joint. You can easily 
palpate, or feel with your fingers, the entire length of your clavicle. 

Axial and Appendicular Skeletons 


Vertebral 
column 


Pelvic 
girdle 
(hip bones) 


w 
i 


Anterior view 


Skull 


Cranial portion 


Facial portion 


Pectoral (shoulder) girdle 


Clavicle 
Scapula 
Thoracic cage 
Sternum 
Ribs 
Upper limb Vertebral 
Humerus column 
vine Pelvic 
Radius girdle 
Carpals (hip bones) 


Metacarpals 


A Ny 


Phalanges 


Lower limb 
Femur 
Patella 
| Key 
Tee © Axial skeleton 
Fibula | Appendicular 
skeleton 

Tarsals 
Metatarsals | 
Phalanges Hy) 


Posterior view 


The axial skeleton forms the central axis of the body and consists of 
the skull, vertebral column, and thoracic cage. The appendicular 
skeleton consists of the pectoral and pelvic girdles, the limb bones, 
and the bones of the hands and feet. 


Pectoral Girdle 


Coracoclavicular Costoclavicular 


ligament ligament 
Clavicle Clavicle 


Acromioclavicular 
joint 


Anterior view of pectoral girdle Posterior view of pectoral girdle 


Posterior 
Lateral 


Anterior 


Superior view of clavicle 


Acromial end Sternal end 


Anterior 


Posterior 


Inferior view of clavicle 


The pectoral girdle consists of the clavicle and the 
scapula, which serve to attach the upper limb to 
the sternum of the axial skeleton. 


The scapula (shoulder blade) lies on the posterior aspect of the shoulder. It 
is supported by the clavicle, which also articulates with the humerus (arm 
bone) to form the shoulder joint. The scapula is a flat, triangular-shaped 


bone with a prominent ridge running across its posterior surface. This ridge 
extends out laterally, where it forms the bony tip of the shoulder and joins 
with the lateral end of the clavicle. By following along the clavicle, you can 
palpate out to the bony tip of the shoulder, and from there, you can move 
back across your posterior shoulder to follow the ridge of the scapula. Move 
your shoulder around and feel how the clavicle and scapula move together 
as a unit. Both of these bones serve as important attachment sites for 
muscles that aid with movements of the shoulder and arm. 


The right and left pectoral girdles are not joined to each other, allowing 
each to operate independently. In addition, the clavicle of each pectoral 
girdle is anchored to the axial skeleton by a single, highly mobile joint. 
This allows for the extensive mobility of the entire pectoral girdle, which in 
turn enhances movements of the shoulder and upper limb. 


Clavicle 


The clavicle is the only long bone that lies in a horizontal position in the 
body (see [link]). The clavicle has several important functions. First, 
anchored by muscles from above, it serves as a strut that extends laterally to 
support the scapula. This in turn holds the shoulder joint superiorly and 
laterally from the body trunk, allowing for maximal freedom of motion for 
the upper limb. The clavicle also transmits forces acting on the upper limb 
to the sternum and axial skeleton. Finally, it serves to protect the underlying 
nerves and blood vessels as they pass between the trunk of the body and the 
upper limb. 


The clavicle has three regions: the medial end, the lateral end, and the shaft. 
The medial end, known as the sternal end of the clavicle, has a triangular 
shape and articulates with the manubrium portion of the sternum. This 
forms the sternoclavicular joint, which is the only bony articulation 
between the pectoral girdle of the upper limb and the axial skeleton. This 
joint allows considerable mobility, enabling the clavicle and scapula to 
move in upward/downward and anterior/posterior directions during 
shoulder movements. The sternoclavicular joint is indirectly supported by 
the costoclavicular ligament (costo- = “rib”), which spans the sternal end 
of the clavicle and the underlying first rib. The lateral or acromial end of 


the clavicle articulates with the acromion of the scapula, the portion of the 
scapula that forms the bony tip of the shoulder. There are some sex 
differences in the morphology of the clavicle. In women, the clavicle tends 
to be shorter, thinner, and less curved. In men, the clavicle is heavier and 
longer, and has a greater curvature and rougher surfaces where muscles 
attach, features that are more pronounced in manual workers. 


The clavicle is the most commonly fractured bone in the body. Such breaks 
often occur because of the force exerted on the clavicle when a person falls 
onto his or her outstretched arms, or when the lateral shoulder receives a 
strong blow. Because the sternoclavicular joint is strong and rarely 
dislocated, excessive force results in the breaking of the clavicle, usually 
between the middle and lateral portions of the bone. If the fracture is 
complete, the shoulder and lateral clavicle fragment will drop due to the 
weight of the upper limb, causing the person to support the sagging limb 
with their other hand. Muscles acting across the shoulder will also pull the 
shoulder and lateral clavicle anteriorly and medially, causing the clavicle 
fragments to override. The clavicle overlies many important blood vessels 
and nerves for the upper limb, but fortunately, due to the anterior 
displacement of a broken clavicle, these structures are rarely affected when 
the clavicle is fractured. 


Scapula 


The scapula is also part of the pectoral girdle and thus plays an important 
role in anchoring the upper limb to the body. The scapula is located on the 
posterior side of the shoulder. It is surrounded by muscles on both its 
anterior (deep) and posterior (superficial) sides, and thus does not articulate 
with the ribs of the thoracic cage. 


The scapula has several important landmarks ([link]). The three margins or 
borders of the scapula, named for their positions within the body, are the 
superior border of the scapula, the medial border of the scapula, and 
the lateral border of the scapula. The suprascapular notch is located 
lateral to the midpoint of the superior border. The corners of the triangular 
scapula, at either end of the medial border, are the superior angle of the 
scapula, located between the medial and superior borders, and the inferior 


angle of the scapula, located between the medial and lateral borders. The 
inferior angle is the most inferior portion of the scapula, and is particularly 
important because it serves as the attachment point for several powerful 
muscles involved in shoulder and upper limb movements. The remaining 
corner of the scapula, between the superior and lateral borders, is the 
location of the glenoid cavity (glenoid fossa). This shallow depression 
articulates with the humerus bone of the arm to form the glenohumeral 
joint (shoulder joint). The small bony bumps located immediately above 
and below the glenoid cavity are the supraglenoid tubercle and the 
infraglenoid tubercle, respectively. These provide attachments for muscles 
of the arm. 

Scapula 


oS Pectoral girdle: 


Acromion Suprascapular Superior border Coracoid process 


Suprascapular notch 


Coracoid F 
process Acromion 
Glenoid 


. Glenoid 
cavity 


cavity 


Superior 
Sy ee angle ee - 
¥ _~—=——_> Supraspinous l-—~\ b 
. \ fossa Te 
; 
\ - ] fae 


ea la 
foss: 


\ ee 
fossa 
\ | ey 
Lateral border \7 Medial border 


=4_—_——_ inferior angle——— 


Lateral border 


Right scapula, anterior aspect Right scapula, posterior aspect 


The isolated scapula is shown here from its 
anterior (deep) side and its posterior (superficial) 
side. 


The scapula also has two prominent projections. Toward the lateral end of 
the superior border, between the suprascapular notch and glenoid cavity, is 
the hook-like coracoid process (coracoid = “shaped like a crow’s beak”). 
This process projects anteriorly and curves laterally. At the shoulder, the 
coracoid process is located inferior to the lateral end of the clavicle. It is 


anchored to the clavicle by a strong ligament, and serves as the attachment 
site for muscles of the anterior chest and arm. On the posterior aspect, the 
spine of the scapula is a long and prominent ridge that runs across its 
upper portion. Extending laterally from the spine is a flattened and 
expanded region called the acromion or acromial process. The acromion 
forms the bony tip of the superior shoulder region and articulates with the 
lateral end of the clavicle, forming the acromioclavicular joint (see [link]). 
Together, the clavicle, acromion, and spine of the scapula form a V-shaped 
bony line that provides for the attachment of neck and back muscles that act 
on the shoulder, as well as muscles that pass across the shoulder joint to act 
on the arm. 


The scapula has three depressions, each of which is called a fossa (plural = 
fossae). Two of these are found on the posterior scapula, above and below 
the scapular spine. Superior to the spine is the narrow supraspinous fossa, 
and inferior to the spine is the broad infraspinous fossa. The anterior 
(deep) surface of the scapula forms the broad subscapular fossa. All of 
these fossae provide large surface areas for the attachment of muscles that 
cross the shoulder joint to act on the humerus. 


The acromioclavicular joint transmits forces from the upper limb to the 
clavicle. The ligaments around this joint are relatively weak. A hard fall 
onto the elbow or outstretched hand can stretch or tear the 
acromioclavicular ligaments, resulting in a moderate injury to the joint. 
However, the primary support for the acromioclavicular joint comes from a 
very strong ligament called the coracoclavicular ligament (see [link]). 
This connective tissue band anchors the coracoid process of the scapula to 
the inferior surface of the acromial end of the clavicle and thus provides 
important indirect support for the acromioclavicular joint. Following a 
strong blow to the lateral shoulder, such as when a hockey player is driven 
into the boards, a complete dislocation of the acromioclavicular joint can 
result. In this case, the acromion is thrust under the acromial end of the 
clavicle, resulting in ruptures of both the acromioclavicular and 
coracoclavicular ligaments. The scapula then separates from the clavicle, 
with the weight of the upper limb pulling the shoulder downward. This 
dislocation injury of the acromioclavicular joint is known as a “shoulder 


separation” and is common in contact sports such as hockey, football, or 
martial arts. 


Chapter Review 


The pectoral girdle, consisting of the clavicle and the scapula, attaches each 
upper limb to the axial skeleton. The clavicle is an anterior bone whose 
sternal end articulates with the manubrium of the sternum at the 
sternoclavicular joint. The sternal end is also anchored to the first rib by the 
costoclavicular ligament. The acromial end of the clavicle articulates with 
the acromion of the scapula at the acromioclavicular joint. This end is also 
anchored to the coracoid process of the scapula by the coracoclavicular 
ligament, which provides indirect support for the acromioclavicular joint. 
The clavicle supports the scapula, transmits the weight and forces from the 
upper limb to the body trunk, and protects the underlying nerves and blood 
vessels. 


The scapula lies on the posterior aspect of the pectoral girdle. It mediates 
the attachment of the upper limb to the clavicle, and contributes to the 
formation of the glenohumeral (shoulder) joint. This triangular bone has 
three sides called the medial, lateral, and superior borders. The 
suprascapular notch is located on the superior border. The scapula also has 
three corners, two of which are the superior and inferior angles. The third 
corner is occupied by the glenoid cavity. Posteriorly, the spine separates the 
supraspinous and infraspinous fossae, and then extends laterally as the 
acromion. The subscapular fossa is located on the anterior surface of the 
scapula. The coracoid process projects anteriorly, passing inferior to the 
lateral end of the clavicle. 


Review Questions 


Exercise: 


Problem: Which part of the clavicle articulates with the manubrium? 


a. shaft 


b. sternal end 
c. acromial end 
d. coracoid process 


Solution: 


B 


Exercise: 


Problem:A shoulder separation results from injury to the 


a. glenohumeral joint 

b. costoclavicular joint 

c. acromioclavicular joint 
d. sternoclavicular joint 


Solution: 


C 
Exercise: 


Problem: 


Which feature lies between the spine and superior border of the 
scapula? 


a. suprascapular notch 
b. glenoid cavity 

c. superior angle 

d. supraspinous fossa 


Solution: 


D 


Exercise: 


Problem: What structure is an extension of the spine of the scapula? 


a. acromion 

b. coracoid process 

c. supraglenoid tubercle 
d. glenoid cavity 


Solution: 


A 
Exercise: 


Problem: 


Name the short, hook-like bony process of the scapula that projects 
anteriorly. 


a. acromial process 
b. clavicle 

c. coracoid process 
d. glenoid fossa 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: 


Describe the shape and palpable line formed by the clavicle and 
scapula. 


Solution: 


The clavicle extends laterally across the anterior shoulder and can be 
palpated along its entire length. At its lateral end, the clavicle 
articulates with the acromion of the scapula, which forms the bony tip 
of the shoulder. The acromion is continuous with the spine of the 
scapula, which can be palpated medially and posteriorly along its 
length. Together, the clavicle, acromion, and spine of the scapula form 
a V-shaped line that serves as an important area for muscle attachment. 


Exercise: 


Problem: 


Discuss two possible injuries of the pectoral girdle that may occur 
following a strong blow to the shoulder or a hard fall onto an 
outstretched hand. 


Solution: 


A blow to the shoulder or falling onto an outstretched hand passes 
strong forces through the scapula to the clavicle and sternum. A hard 
fall may thus cause a fracture of the clavicle (broken collarbone) or 
may injure the ligaments of the acromioclavicular joint. In a severe 
case, the coracoclavicular ligament may also rupture, resulting in 
complete dislocation of the acromioclavicular joint (a “shoulder 
separation”). 


Glossary 


acromial end of the clavicle 
lateral end of the clavicle that articulates with the acromion of the 
scapula 


acromial process 
acromion of the scapula 


acromioclavicular joint 
articulation between the acromion of the scapula and the acromial end 
of the clavicle 


acromion 
flattened bony process that extends laterally from the scapular spine to 
form the bony tip of the shoulder 


clavicle 
collarbone; elongated bone that articulates with the manubrium of the 
sternum medially and the acromion of the scapula laterally 


coracoclavicular ligament 
strong band of connective tissue that anchors the coracoid process of 
the scapula to the lateral clavicle; provides important indirect support 
for the acromioclavicular joint 


coracoid process 
short, hook-like process that projects anteriorly and laterally from the 
superior margin of the scapula 


costoclavicular ligament 
band of connective tissue that unites the medial clavicle with the first 
rib 

fossa 


(plural = fossae) shallow depression on the surface of a bone 


glenohumeral joint 
shoulder joint; formed by the articulation between the glenoid cavity 
of the scapula and the head of the humerus 


glenoid cavity 
(also, glenoid fossa) shallow depression located on the lateral scapula, 


between the superior and lateral borders 


inferior angle of the scapula 


inferior corner of the scapula located where the medial and lateral 
borders meet 


infraglenoid tubercle 
small bump or roughened area located on the lateral border of the 
scapula, near the inferior margin of the glenoid cavity 


infraspinous fossa 
broad depression located on the posterior scapula, inferior to the spine 


lateral border of the scapula 
diagonally oriented lateral margin of the scapula 


medial border of the scapula 
elongated, medial margin of the scapula 


pectoral girdle 
shoulder girdle; the set of bones, consisting of the scapula and clavicle, 
which attaches each upper limb to the axial skeleton 


scapula 
shoulder blade bone located on the posterior side of the shoulder 


spine of the scapula 
prominent ridge passing mediolaterally across the upper portion of the 
posterior scapular surface 


sternal end of the clavicle 
medial end of the clavicle that articulates with the manubrium of the 
sternum 


sternoclavicular joint 
articulation between the manubrium of the sternum and the sternal end 
of the clavicle; forms the only bony attachment between the pectoral 
girdle of the upper limb and the axial skeleton 


subscapular fossa 
broad depression located on the anterior (deep) surface of the scapula 


superior angle of the scapula 
comer of the scapula between the superior and medial borders of the 
scapula 


superior border of the scapula 
superior margin of the scapula 


supraglenoid tubercle 
small bump located at the superior margin of the glenoid cavity 


suprascapular notch 
small notch located along the superior border of the scapula, medial to 
the coracoid process 


supraspinous fossa 
narrow depression located on the posterior scapula, superior to the 
spine 


Bones of the Upper Limb 
By the end of this section, you will be able to: 


e Identify the divisions of the upper limb and describe the bones in each 
region 

e List the bones and bony landmarks that articulate at each joint of the 
upper limb 


The upper limb is divided into three regions. These consist of the arm, 
located between the shoulder and elbow joints; the forearm, which is 
between the elbow and wrist joints; and the hand, which is located distal to 
the wrist. There are 30 bones in each upper limb (see [link]). The humerus 
is the single bone of the upper arm, and the ulna (medially) and the radius 
(laterally) are the paired bones of the forearm. The base of the hand 
contains eight bones, each called a carpal bone, and the palm of the hand is 
formed by five bones, each called a metacarpal bone. The fingers and 
thumb contain a total of 14 bones, each of which is a phalanx bone of the 
hand. 


Humerus 


The humerus is the single bone of the upper arm region ([link]). At its 
proximal end is the head of the humerus. This is the large, round, smooth 
region that faces medially. The head articulates with the glenoid cavity of 
the scapula to form the glenohumeral (shoulder) joint. The margin of the 
smooth area of the head is the anatomical neck of the humerus. Located on 
the lateral side of the proximal humerus is an expanded bony area called the 
greater tubercle. The smaller lesser tubercle of the humerus is found on 
the anterior aspect of the humerus. Both the greater and lesser tubercles 
serve as attachment sites for muscles that act across the shoulder joint. 
Passing between the greater and lesser tubercles is the narrow 
intertubercular groove (sulcus), which is also known as the bicipital 
groove because it provides passage for a tendon of the biceps brachii 
muscle. The surgical neck is located at the base of the expanded, proximal 
end of the humerus, where it joins the narrow shaft of the humerus. The 
surgical neck is a common site of arm fractures. The deltoid tuberosity is a 
roughened, V-shaped region located on the lateral side in the middle of the 


humerus shaft. As its name indicates, it is the site of attachment for the 
deltoid muscle. 
Humerus and Elbow Joint 


Humerus 


Anatomical 
Greater (Oy, Gin Greater 
tubercle 1 \ ) Head tubercle 


Lesser 
tubercle 


Intertubercular 
groove (sulcus) 


Deltoid 
| tuberosity 


| Body (shaft) 
Lateral 
supracondylar 
ridge 


Radial 
fossa 


Olecranon fossa Lateral 


Coronoid fossa epicondyle 
Olecranon 
of ulna 


Head of 
radius 


Capitulum —+~—— Medial 


Head of epicondyle 


radius Trochlea 


Coronoid process 
of ulna 


Anterior view Posterior view 


The humerus is the single bone of the 
upper arm region. It articulates with 
the radius and ulna bones of the 
forearm to form the elbow joint. 


Distally, the humerus becomes flattened. The prominent bony projection on 
the medial side is the medial epicondyle of the humerus. The much 
smaller lateral epicondyle of the humerus is found on the lateral side of 
the distal humerus. The roughened ridge of bone above the lateral 
epicondyle is the lateral supracondylar ridge. All of these areas are 
attachment points for muscles that act on the forearm, wrist, and hand. The 
powerful grasping muscles of the anterior forearm arise from the medial 


epicondyle, which is thus larger and more robust than the lateral epicondyle 
that gives rise to the weaker posterior forearm muscles. 


The distal end of the humerus has two articulation areas, which join the 
ulna and radius bones of the forearm to form the elbow joint. The more 
medial of these areas is the trochlea, a spindle- or pulley-shaped region 
(trochlea = “pulley”), which articulates with the ulna bone. Immediately 
lateral to the trochlea is the capitulum (“small head”), a knob-like structure 
located on the anterior surface of the distal humerus. The capitulum 
articulates with the radius bone of the forearm. Just above these bony areas 
are two small depressions. These spaces accommodate the forearm bones 
when the elbow is fully bent (flexed). Superior to the trochlea is the 
coronoid fossa, which receives the coronoid process of the ulna, and above 
the capitulum is the radial fossa, which receives the head of the radius 
when the elbow is flexed. Similarly, the posterior humerus has the 
olecranon fossa, a larger depression that receives the olecranon process of 
the ulna when the forearm is fully extended. 


Ulna 


The ulna is the medial bone of the forearm. It runs parallel to the radius, 
which is the lateral bone of the forearm ([link]). The proximal end of the 
ulna resembles a crescent wrench with its large, C-shaped trochlear notch. 
This region articulates with the trochlea of the humerus as part of the elbow 
joint. The inferior margin of the trochlear notch is formed by a prominent 
lip of bone called the coronoid process of the ulna. Just below this on the 
anterior ulna is a roughened area called the ulnar tuberosity. To the lateral 
side and slightly inferior to the trochlear notch is a small, smooth area 
called the radial notch of the ulna. This area is the site of articulation 
between the proximal radius and the ulna, forming the proximal 
radioulnar joint. The posterior and superior portions of the proximal ulna 
make up the olecranon process, which forms the bony tip of the elbow. 
Ulna and Radius 


er a ie Olecranon 
Radial notch on process 
of the ulna 1S 
Trochlear Head of 
Head of notch radius 
radius 
Neck of Coronoid 
di process Neck of 
radius radius 
Radial Proximal 
tuberosity radioulnar 
} joint 
| Interosseous 
\\ | membrane 
\ \ \ Uina 
Radius S 
Ulnar notch 
of the radius 
: Head of ulna Radius 
Distal 
radioulnar 


joint 


Styloid process 
of ulna 


Styloid process 
of radius 


The ulna is located on the medial side 
of the forearm, and the radius is on the 
lateral side. These bones are attached 
to each other by an interosseous 
membrane. 


More distal is the shaft of the ulna. The lateral side of the shaft forms a 
ridge called the interosseous border of the ulna. This is the line of 
attachment for the interosseous membrane of the forearm, a sheet of 
dense connective tissue that unites the ulna and radius bones. The small, 
rounded area that forms the distal end is the head of the ulna. Projecting 
from the posterior side of the ulnar head is the styloid process of the ulna, 
a short bony projection. This serves as an attachment point for a connective 
tissue structure that unites the distal ends of the ulna and radius. 


In the anatomical position, with the elbow fully extended and the palms 
facing forward, the arm and forearm do not form a straight line. Instead, the 
forearm deviates laterally by 5-15 degrees from the line of the arm. This 
deviation is called the carrying angle. It allows the forearm and hand to 
swing freely or to carry an object without hitting the hip. The carrying angle 
is larger in females to accommodate their wider pelvis. 


Radius 


The radius runs parallel to the ulna, on the lateral (thumb) side of the 
forearm (see [link]). The head of the radius is a disc-shaped structure that 
forms the proximal end. The small depression on the surface of the head 
articulates with the capitulum of the humerus as part of the elbow joint, 
whereas the smooth, outer margin of the head articulates with the radial 
notch of the ulna at the proximal radioulnar joint. The neck of the radius is 
the narrowed region immediately below the expanded head. Inferior to this 
point on the medial side is the radial tuberosity, an oval-shaped, bony 
protuberance that serves as a muscle attachment point. The shaft of the 
radius is slightly curved and has a small ridge along its medial side. This 
ridge forms the interosseous border of the radius, which, like the similar 
border of the ulna, is the line of attachment for the interosseous membrane 
that unites the two forearm bones. The distal end of the radius has a smooth 
surface for articulation with two carpal bones to form the radiocarpal joint 
or wrist joint ({link] and [link]). On the medial side of the distal radius is 
the ulnar notch of the radius. This shallow depression articulates with the 
head of the ulna, which together form the distal radioulnar joint. The 
lateral end of the radius has a pointed projection called the styloid process 
of the radius. This provides attachment for ligaments that support the 
lateral side of the wrist joint. Compared to the styloid process of the ulna, 
the styloid process of the radius projects more distally, thereby limiting the 
range of movement for lateral deviations of the hand at the wrist joint. 


Note: 


Watch this video to see how fractures of the distal radius bone can affect 
the wrist joint. Explain the problems that may occur if a fracture of the 
distal radius involves the joint surface of the radiocarpal joint of the wrist. 


Carpal Bones 


The wrist and base of the hand are formed by a series of eight small carpal 
bones (see [link]). The carpal bones are arranged in two rows, forming a 
proximal row of four carpal bones and a distal row of four carpal bones. 
The bones in the proximal row, running from the lateral (thumb) side to the 
medial side, are the scaphoid (“boat-shaped”), lunate (“moon-shaped”), 
triquetrum (“three-cornered”), and pisiform (“pea-shaped”) bones. The 
small, rounded pisiform bone articulates with the anterior surface of the 
triquetrum bone. The pisiform thus projects anteriorly, where it forms the 
bony bump that can be felt at the medial base of your hand. The distal 
bones (lateral to medial) are the trapezium (“table”), trapezoid 
(“resembles a table”), capitate (“head-shaped”), and hamate (“hooked 
bone”) bones. The hamate bone is characterized by a prominent bony 
extension on its anterior side called the hook of the hamate bone. 


A helpful mnemonic for remembering the arrangement of the carpal bones 
is “So Long To Pinky, Here Comes The Thumb.” This mnemonic starts on 
the lateral side and names the proximal bones from lateral to medial 
(scaphoid, lunate, triquetrum, pisiform), then makes a U-turn to name the 
distal bones from medial to lateral (hamate, capitate, trapezoid, trapezium). 
Thus, it starts and finishes on the lateral side. 

Bones of the Wrist and Hand 


| Carpals 


S i\\ \ ~;Metacarpals 


yyy ib Phalanges 


Middle finger 
Index finger Ring finger 
4 Distal 
Little finger . 
Miceie Phalanges 
Thumb f 
(pollex) 
Phalanges: /f ie 
Distal Head —¥~q — 
Proximal a Shaft \ 
Base 
Head 
Metacarpals v) 
(1-5) J Shaft 
Carpals: \ y . : 
Trapezium - a on Va j Base 
Trapezoid t ) — ; _=—_ : 
f - Capitate : Carpals: 
Scaphoid x ‘< Pisiform ) : Trapezium 
VW y> Triquetrum p =: Trapeziod 
" \ f Lunate - Scaphoid 
7 Ulna 
| Radius 


Anterior view Posterior view 


The eight carpal bones form the base of the hand. 
These are arranged into proximal and distal rows of 
four bones each. The metacarpal bones form the palm 
of the hand. The thumb and fingers consist of the 
phalanx bones. 


The carpal bones form the base of the hand. This can be seen in the 
radiograph (X-ray image) of the hand that shows the relationships of the 
hand bones to the skin creases of the hand (see [link]). Within the carpal 
bones, the four proximal bones are united to each other by ligaments to 
form a unit. Only three of these bones, the scaphoid, lunate, and triquetrum, 
contribute to the radiocarpal joint. The scaphoid and lunate bones articulate 
directly with the distal end of the radius, whereas the triquetrum bone 
articulates with a fibrocartilaginous pad that spans the radius and styloid 
process of the ulna. The distal end of the ulna thus does not directly 
articulate with any of the carpal bones. 


The four distal carpal bones are also held together as a group by ligaments. 
The proximal and distal rows of carpal bones articulate with each other to 
form the midcarpal joint (see [link]). Together, the radiocarpal and 
midcarpal joints are responsible for all movements of the hand at the wrist. 
The distal carpal bones also articulate with the metacarpal bones of the 
hand. 

Bones of the Hand 


Metacarpophalangeal 


joints Interphalangeal joints 


Carpometacarpal 
joints 


Midcarpal joint 
Radiocarpal (wrist) 


fs : 


This radiograph shows the position of the bones within 
the hand. Note the carpal bones that form the base of 
the hand. (credit: modification of work by Trace 
Meek) 


In the articulated hand, the carpal bones form a U-shaped grouping. A 
strong ligament called the flexor retinaculum spans the top of this U- 
shaped area to maintain this grouping of the carpal bones. The flexor 
retinaculum is attached laterally to the trapezium and scaphoid bones, and 
medially to the hamate and pisiform bones. Together, the carpal bones and 
the flexor retinaculum form a passageway called the carpal tunnel, with 
the carpal bones forming the walls and floor, and the flexor retinaculum 
forming the roof of this space ({link]). The tendons of nine muscles of the 


anterior forearm and an important nerve pass through this narrow tunnel to 
enter the hand. Overuse of the muscle tendons or wrist injury can produce 
inflammation and swelling within this space. This produces compression of 
the nerve, resulting in carpal tunnel syndrome, which is characterized by 
pain or numbness, and muscle weakness in those areas of the hand supplied 
by this nerve. 

Carpal Tunnel 


Carpal 
tunnel 


Muscle 
tendons 


— Flexor 


\ 
rpal bones: — i 
Carpal bones retinaculum 


Hamate 
Trapezoid 
Trapezium 
Capitate 


Nerve 


The carpal tunnel is the passageway 
by which nine muscle tendons and a 
major nerve enter the hand from the 
anterior forearm. The walls and floor 
of the carpal tunnel are formed by the 
U-shaped grouping of the carpal 
bones, and the roof is formed by the 
flexor retinaculum, a strong ligament 
that anteriorly unites the bones. 


Metacarpal Bones 


The palm of the hand contains five elongated metacarpal bones. These 
bones lie between the carpal bones of the wrist and the bones of the fingers 
and thumb (see [link]). The proximal end of each metacarpal bone 
articulates with one of the distal carpal bones. Each of these articulations is 
a Carpometacarpal joint (see [link]). The expanded distal end of each 
metacarpal bone articulates at the metacarpophalangeal joint with the 
proximal phalanx bone of the thumb or one of the fingers. The distal end 
also forms the knuckles of the hand, at the base of the fingers. The 
metacarpal bones are numbered 1-5, beginning at the thumb. 


The first metacarpal bone, at the base of the thumb, is separated from the 
other metacarpal bones. This allows it a freedom of motion that is 
independent of the other metacarpal bones, which is very important for 
thumb mobility. The remaining metacarpal bones are united together to 
form the palm of the hand. The second and third metacarpal bones are 
firmly anchored in place and are immobile. However, the fourth and fifth 
metacarpal bones have limited anterior-posterior mobility, a motion that is 
greater for the fifth bone. This mobility is important during power gripping 
with the hand ([link]). The anterior movement of these bones, particularly 
the fifth metacarpal bone, increases the strength of contact for the medial 
hand during gripping actions. 

Hand During Gripping 


(a) Loosely held (b) Firmly gripped 


During tight gripping—compare (b) to (a)—the fourth 
and, particularly, the fifth metatarsal bones are pulled 
anteriorly. This increases the contact between the 
object and the medial side of the hand, thus improving 
the firmness of the grip. 


Phalanx Bones 


The fingers and thumb contain 14 bones, each of which is called a phalanx 
bone (plural = phalanges), named after the ancient Greek phalanx (a 
rectangular block of soldiers). The thumb (pollex) is digit number 1 and has 
two phalanges, a proximal phalanx, and a distal phalanx bone (see [link]). 
Digits 2 (index finger) through 5 (little finger) have three phalanges each, 
called the proximal, middle, and distal phalanx bones. An interphalangeal 
joint is one of the articulations between adjacent phalanges of the digits 
(see [Link]). 


Note: 


ORs sao 
at ; 


— . 
=——: 
wm OPENSTAX COLLEGE 


Visit this site to explore the bones and joints of the hand. What are the 
three arches of the hand, and what is the importance of these during the 
sripping of an object? 


Note: 

Disorders of the... 

Appendicular System: Fractures of Upper Limb Bones 

Due to our constant use of the hands and the rest of our upper limbs, an 
injury to any of these areas will cause a significant loss of functional 
ability. Many fractures result from a hard fall onto an outstretched hand. 
The resulting transmission of force up the limb may result in a fracture of 
the humerus, radius, or scaphoid bones. These injuries are especially 
common in elderly people whose bones are weakened due to osteoporosis. 


Falls onto the hand or elbow, or direct blows to the arm, can result in 
fractures of the humerus ((link]). Following a fall, fractures at the surgical 
neck, the region at which the expanded proximal end of the humerus joins 
with the shaft, can result in an impacted fracture, in which the distal 
portion of the humerus is driven into the proximal portion. Falls or blows 
to the arm can also produce transverse or spiral fractures of the humeral 
shaft. 

In children, a fall onto the tip of the elbow frequently results in a distal 
humerus fracture. In these, the olecranon of the ulna is driven upward, 
resulting in a fracture across the distal humerus, above both epicondyles 
(supracondylar fracture), or a fracture between the epicondyles, thus 
separating one or both of the epicondyles from the body of the humerus 
(intercondylar fracture). With these injuries, the immediate concern is 
possible compression of the artery to the forearm due to swelling of the 
surrounding tissues. If compression occurs, the resulting ischemia (lack of 
oxygen) due to reduced blood flow can quickly produce irreparable 
damage to the forearm muscles. In addition, four major nerves for shoulder 
and upper limb muscles are closely associated with different regions of the 
humerus, and thus, humeral fractures may also damage these nerves. 
Another frequent injury following a fall onto an outstretched hand is a 
Colles fracture (“col-lees”’) of the distal radius (see [link]). This involves a 
complete transverse fracture across the distal radius that drives the 
separated distal fragment of the radius posteriorly and superiorly. This 
injury results in a characteristic “dinner fork” bend of the forearm just 
above the wrist due to the posterior displacement of the hand. This is the 
most frequent forearm fracture and is a common injury in persons over the 
age of 50, particularly in older women with osteoporosis. It also commonly 
occurs following a high-speed fall onto the hand during activities such as 
snowboarding or skating. 

The most commonly fractured carpal bone is the scaphoid, often resulting 
from a fall onto the hand. Deep pain at the lateral wrist may yield an initial 
diagnosis of a wrist sprain, but a radiograph taken several weeks after the 
injury, after tissue swelling has subsided, will reveal the fracture. Due to 
the poor blood supply to the scaphoid bone, healing will be slow and there 
is the danger of bone necrosis and subsequent degenerative joint disease of 
the wrist. 


Fractures of the Humerus and Radius 


Surgical neck fracture 


Transverse humeral fracture 


Fractures of the Humerus 


Supracondylar fracture 


KL 
i 
Eas 


Normal 


Normal 


Colles Fracture of the Distal Radius 


Falls or direct blows can result in fractures of 
the surgical neck or shaft of the humerus. Falls 
onto the elbow can fracture the distal humerus. 

A Colles fracture of the distal radius is the 
most common forearm fracture. 


Note: 


[=] 


[eae 


1 


openstax COLLEGE” 


Watch this video to learn about a Colles fracture, a break of the distal 
radius, usually caused by falling onto an outstretched hand. When would 
surgery be required and how would the fracture be repaired in this case? 


Chapter Review 


Each upper limb is divided into three regions and contains a total of 30 
bones. The upper arm is the region located between the shoulder and elbow 
joints. This area contains the humerus. The proximal humerus consists of 
the head, which articulates with the scapula at the glenohumeral joint, the 
greater and lesser tubercles separated by the intertubercular (bicipital) 
groove, and the anatomical and surgical necks. The humeral shaft has the 
roughened area of the deltoid tuberosity on its lateral side. The distal 
humerus is flattened, forming a lateral supracondylar ridge that terminates 
at the small lateral epicondyle. The medial side of the distal humerus has 
the large, medial epicondyle. The articulating surfaces of the distal humerus 
consist of the trochlea medially and the capitulum laterally. Depressions on 
the humerus that accommodate the forearm bones during bending (flexing) 
and straightening (extending) of the elbow include the coronoid fossa, the 
radial fossa, and the olecranon fossa. 


The forearm is the region of the upper limb located between the elbow and 
wrist joints. This region contains two bones, the ulna medially and the 
radius on the lateral (thumb) side. The elbow joint is formed by the 
articulation between the trochlea of the humerus and the trochlear notch of 
the ulna, plus the articulation between the capitulum of the humerus and the 
head of the radius. The proximal radioulnar joint is the articulation between 


the head of the radius and the radial notch of the ulna. The proximal ulna 
also has the olecranon process, forming an expanded posterior region, and 
the coronoid process and ulnar tuberosity on its anterior aspect. On the 
proximal radius, the narrowed region below the head is the neck; distal to 
this is the radial tuberosity. The shaft portions of both the ulna and radius 
have an interosseous border, whereas the distal ends of each bone have a 
pointed styloid process. The distal radioulnar joint is found between the 
head of the ulna and the ulnar notch of the radius. The distal end of the 
radius articulates with the proximal carpal bones, but the ulna does not. 


The base of the hand is formed by eight carpal bones. The carpal bones are 
united into two rows of bones. The proximal row contains (from lateral to 
medial) the scaphoid, lunate, triquetrum, and pisiform bones. The scaphoid, 
lunate, and triquetrum bones contribute to the formation of the radiocarpal 
joint. The distal row of carpal bones contains (from medial to lateral) the 
hamate, capitate, trapezoid, and trapezium bones (“So Long To Pinky, Here 
Comes The Thumb”). The anterior hamate has a prominent bony hook. The 
proximal and distal carpal rows articulate with each other at the midcarpal 
joint. The carpal bones, together with the flexor retinaculum, also form the 
carpal tunnel of the wrist. 


The five metacarpal bones form the palm of the hand. The metacarpal bones 
are numbered 1-5, starting with the thumb side. The first metacarpal bone 
is freely mobile, but the other bones are united as a group. The digits are 
also numbered 1—5, with the thumb being number 1. The fingers and thumb 
contain a total of 14 phalanges (phalanx bones). The thumb contains a 
proximal and a distal phalanx, whereas the remaining digits each contain 
proximal, middle, and distal phalanges. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to see how fractures of the distal radius bone can 
affect the wrist joint. Explain the problems that may occur if a fracture 
of the distal radius involves the joint surface of the radiocarpal joint of 
the wrist. 


Solution: 


A fracture through the joint surface of the distal radius may make the 
articulating surface of the radius rough or jagged. This can then cause 
painful movements involving this joint and the early development of 
arthritis. Surgery can return the joint surface to its original smoothness, 
thus allowing for the return of normal function. 


Exercise: 


Problem: 


Visit this site to explore the bones and joints of the hand. What are the 
three arches of the hand, and what is the importance of these during 
the gripping of an object? 


Solution: 


The hand has a proximal transverse arch, a distal transverse arch, and a 
longitudinal arch. These allow the hand to conform to objects being 
held. These arches maximize the amount of surface contact between 
the hand and object, which enhances stability and increases sensory 
input. 


Exercise: 
Problem: 
Watch this video to learn about a Colles fracture, a break of the distal 
radius, usually caused by falling onto an outstretched hand. When 


would surgery be required and how would the fracture be repaired in 
this case? 


Solution: 


Surgery may be required if the fracture is unstable, meaning that the 
broken ends of the radius won’t stay in place to allow for proper 
healing. In this case, metal plates and screws can be used to stabilize 
the fractured bone. 


Review Questions 


Exercise: 


Problem:How many bones are there in the upper limbs combined? 


a. 20 
b. 30 
c. 40 
d. 60 


Solution: 


D 
Exercise: 
Problem: 


Which bony landmark is located on the lateral side of the proximal 
humerus? 


a. greater tubercle 
b. trochlea 

c. lateral epicondyle 
d. lesser tubercle 


Solution: 


A 
Exercise: 


Problem: 


Which region of the humerus articulates with the radius as part of the 
elbow joint? 


a. trochlea 

b. styloid process 

c. capitulum 

d. olecranon process 


Solution: 


C 


Exercise: 


Problem: Which is the lateral-most carpal bone of the proximal row? 


a. trapezium 
b. hamate 

c. pisiform 
d. scaphoid 


Solution: 
D 
Exercise: 


Problem: The radius bone 


a. is found on the medial side of the forearm 
b. has a head that articulates with the radial notch of the ulna 
c. does not articulate with any of the carpal bones 


d. has the radial tuberosity located near its distal end 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


Your friend runs out of gas and you have to help push his car. Discuss 
the sequence of bones and joints that convey the forces passing from 
your hand, through your upper limb and your pectoral girdle, and to 
your axial skeleton. 


Solution: 


As you push against the car, forces will pass from the metacarpal 
bones of your hand into the carpal bones at the base of your hand. 
Forces will then pass through the midcarpal and radiocarpal joints into 
the radius and ulna bones of the forearm. These will pass the force 
through the elbow joint into the humerus of the arm, and then through 
the glenohumeral joint into the scapula. The force will travel through 
the acromioclavicular joint into the clavicle, and then through the 
sternoclavicular joint into the sternum, which is part of the axial 
skeleton. 


Exercise: 


Problem: 


Name the bones in the wrist and hand, and describe or sketch out their 
locations and articulations. 


Solution: 


The base of the hand is formed by the eight carpal bones arranged in 
two rows (distal and proximal) of four bones each. The proximal row 
contains (from lateral to medial) the scaphoid, lunate, triquetrum, and 
pisiform bones. The distal row contains (from medial to lateral) the 
hamate, capitate, trapezoid, and trapezium bones. (Use the mnemonic 
“So Long To Pinky, Here Comes The Thumb” to remember this 
sequence). The rows of the proximal and distal carpal bones articulate 
with each other at the midcarpal joint. The palm of the hand contains 
the five metacarpal bones, which are numbered 1-5 starting on the 
thumb side. The proximal ends of the metacarpal bones articulate with 
the distal row of the carpal bones. The distal ends of the metacarpal 
bones articulate with the proximal phalanx bones of the thumb and 
fingers. The thumb (digit 1) has both a proximal and distal phalanx 
bone. The fingers (digits 2—5) all contain proximal, middle, and distal 
phalanges. 


Glossary 


anatomical neck 
line on the humerus located around the outside margin of the humeral 
head 


arm 
region of the upper limb located between the shoulder and elbow 
joints; contains the humerus bone 


bicipital groove 
intertubercular groove; narrow groove located between the greater and 
lesser tubercles of the humerus 


capitate 
from the lateral side, the third of the four distal carpal bones; 
articulates with the scaphoid and lunate proximally, the trapezoid 
laterally, the hamate medially, and primarily with the third metacarpal 
distally 


capitulum 


knob-like bony structure located anteriorly on the lateral, distal end of 
the humerus 


carpal bone 
one of the eight small bones that form the wrist and base of the hand; 
these are grouped as a proximal row consisting of (from lateral to 
medial) the scaphoid, lunate, triquetrum, and pisiform bones, and a 
distal row containing (from lateral to medial) the trapezium, trapezoid, 
capitate, and hamate bones 


carpal tunnel 
passageway between the anterior forearm and hand formed by the 
carpal bones and flexor retinaculum 


Carpometacarpal joint 
articulation between one of the carpal bones in the distal row and a 
metacarpal bone of the hand 


coronoid fossa 
depression on the anterior surface of the humerus above the trochlea; 
this space receives the coronoid process of the ulna when the elbow is 
maximally flexed 


coronoid process of the ulna 
projecting bony lip located on the anterior, proximal ulna; forms the 
inferior margin of the trochlear notch 


deltoid tuberosity 
roughened, V-shaped region located laterally on the mid-shaft of the 
humerus 


distal radioulnar joint 
articulation between the head of the ulna and the ulnar notch of the 
radius 


elbow joint 
joint located between the upper arm and forearm regions of the upper 
limb; formed by the articulations between the trochlea of the humerus 


and the trochlear notch of the ulna, and the capitulum of the humerus 
and the head of the radius 


flexor retinaculum 
strong band of connective tissue at the anterior wrist that spans the top 
of the U-shaped grouping of the carpal bones to form the roof of the 
carpal tunnel 


forearm 
region of the upper limb located between the elbow and wrist joints; 
contains the radius and ulna bones 


greater tubercle 
enlarged prominence located on the lateral side of the proximal 
humerus 


hamate 
from the lateral side, the fourth of the four distal carpal bones; 
articulates with the lunate and triquetrum proximally, the fourth and 
fifth metacarpals distally, and the capitate laterally 


hand 
region of the upper limb distal to the wrist joint 


head of the humerus 
smooth, rounded region on the medial side of the proximal humerus; 
articulates with the glenoid fossa of the scapula to form the 
glenohumeral (shoulder) joint 


head of the radius 
disc-shaped structure that forms the proximal end of the radius; 
articulates with the capitulum of the humerus as part of the elbow 
joint, and with the radial notch of the ulna as part of the proximal 
radioulnar joint 


head of the ulna 
small, rounded distal end of the ulna; articulates with the ulnar notch 
of the distal radius, forming the distal radioulnar joint 


hook of the hamate bone 
bony extension located on the anterior side of the hamate carpal bone 


humerus 
single bone of the upper arm 


interosseous border of the radius 
narrow ridge located on the medial side of the radial shaft; for 
attachment of the interosseous membrane between the ulna and radius 
bones 


interosseous border of the ulna 
narrow ridge located on the lateral side of the ulnar shaft; for 
attachment of the interosseous membrane between the ulna and radius 


interosseous membrane of the forearm 
sheet of dense connective tissue that unites the radius and ulna bones 


interphalangeal joint 
articulation between adjacent phalanx bones of the hand or foot digits 


intertubercular groove (sulcus) 
bicipital groove; narrow groove located between the greater and lesser 
tubercles of the humerus 


lateral epicondyle of the humerus 
small projection located on the lateral side of the distal humerus 


lateral supracondylar ridge 
narrow, bony ridge located along the lateral side of the distal humerus, 
superior to the lateral epicondyle 


lesser tubercle 
small, bony prominence located on anterior side of the proximal 
humerus 


lunate 


from the lateral side, the second of the four proximal carpal bones; 
articulates with the radius proximally, the capitate and hamate distally, 
the scaphoid laterally, and the triquetrum medially 


medial epicondyle of the humerus 
enlarged projection located on the medial side of the distal humerus 


metacarpal bone 
one of the five long bones that form the palm of the hand; numbered 
1—5, starting on the lateral (thumb) side of the hand 


metacarpophalangeal joint 
articulation between the distal end of a metacarpal bone of the hand 
and a proximal phalanx bone of the thumb or a finger 


midcarpal joint 
articulation between the proximal and distal rows of the carpal bones; 
contributes to movements of the hand at the wrist 


neck of the radius 
narrowed region immediately distal to the head of the radius 


olecranon fossa 
large depression located on the posterior side of the distal humerus; 
this space receives the olecranon process of the ulna when the elbow is 
fully extended 


olecranon process 
expanded posterior and superior portions of the proximal ulna; forms 
the bony tip of the elbow 


phalanx bone of the hand 
(plural = phalanges) one of the 14 bones that form the thumb and 
fingers; these include the proximal and distal phalanges of the thumb, 
and the proximal, middle, and distal phalanx bones of the fingers two 
through five 


pisiform 


from the lateral side, the fourth of the four proximal carpal bones; 
articulates with the anterior surface of the triquetrum 


pollex 
(also, thumb) digit 1 of the hand 


proximal radioulnar joint 
articulation formed by the radial notch of the ulna and the head of the 
radius 


radial fossa 
small depression located on the anterior humerus above the capitulum; 
this space receives the head of the radius when the elbow is maximally 
flexed 


radial notch of the ulna 
small, smooth area on the lateral side of the proximal ulna; articulates 
with the head of the radius as part of the proximal radioulnar joint 


radial tuberosity 
oval-shaped, roughened protuberance located on the medial side of the 
proximal radius 


radiocarpal joint 
wrist joint, located between the forearm and hand regions of the upper 
limb; articulation formed proximally by the distal end of the radius and 
the fibrocartilaginous pad that unites the distal radius and ulna bone, 
and distally by the scaphoid, lunate, and triquetrum carpal bones 


radius 
bone located on the lateral side of the forearm 


scaphoid 
from the lateral side, the first of the four proximal carpal bones; 
articulates with the radius proximally, the trapezoid, trapezium, and 
capitate distally, and the lunate medially 


shaft of the humerus 


narrow, elongated, central region of the humerus 


shaft of the radius 
narrow, elongated, central region of the radius 


shaft of the ulna 
narrow, elongated, central region of the ulna 


styloid process of the radius 
pointed projection located on the lateral end of the distal radius 


styloid process of the ulna 
short, bony projection located on the medial end of the distal ulna 


surgical neck 
region of the humerus where the expanded, proximal end joins with 
the narrower shaft 


trapezium 
from the lateral side, the first of the four distal carpal bones; articulates 
with the scaphoid proximally, the first and second metacarpals distally, 
and the trapezoid medially 


trapezoid 
from the lateral side, the second of the four distal carpal bones; 
articulates with the scaphoid proximally, the second metacarpal 
distally, the trapezium laterally, and the capitate medially 


triquetrum 
from the lateral side, the third of the four proximal carpal bones; 
articulates with the lunate laterally, the hamate distally, and has a facet 
for the pisiform 


trochlea 
pulley-shaped region located medially at the distal end of the humerus; 
articulates at the elbow with the trochlear notch of the ulna 


trochlear notch 


large, C-shaped depression located on the anterior side of the proximal 
ulna; articulates at the elbow with the trochlea of the humerus 


ulna 
bone located on the medial side of the forearm 


ulnar notch of the radius 
shallow, smooth area located on the medial side of the distal radius; 
articulates with the head of the ulna at the distal radioulnar joint 


ulnar tuberosity 
roughened area located on the anterior, proximal ulna inferior to the 
coronoid process 


The Pelvic Girdle and Pelvis 
By the end of this section, you will be able to: 


¢ Define the pelvic girdle and describe the bones and ligaments of the 
pelvis 

e Explain the three regions of the hip bone and identify their bony 
landmarks 

e Describe the openings of the pelvis and the boundaries of the greater 
and lesser pelvis 


The pelvic girdle (hip girdle) is formed by a single bone, the hip bone or 
coxal bone (coxal = “hip”), which serves as the attachment point for each 
lower limb. Each hip bone, in turn, is firmly joined to the axial skeleton via 
its attachment to the sacrum of the vertebral column. The right and left hip 
bones also converge anteriorly to attach to each other. The bony pelvis is 
the entire structure formed by the two hip bones, the sacrum, and, attached 
inferiorly to the sacrum, the coccyx ([link]). 


Unlike the bones of the pectoral girdle, which are highly mobile to enhance 
the range of upper limb movements, the bones of the pelvis are strongly 
united to each other to form a largely immobile, weight-bearing structure. 
This is important for stability because it enables the weight of the body to 
be easily transferred laterally from the vertebral column, through the pelvic 
girdle and hip joints, and into either lower limb whenever the other limb is 
not bearing weight. Thus, the immobility of the pelvis provides a strong 
foundation for the upper body as it rests on top of the mobile lower limbs. 
Pelvis 


Sacroiliac joint 


Sacral promonitory 


Sacrum 


Pelvic brim 
Acetabulum 
Coccyx 


Obturator f <i 
ical ieti \ ax Pubic symphysis 
Ischial tuberosity 
ae FH ~~ 


Ischiopubic ramus 


The pelvic girdle is formed by a single hip bone. The 
hip bone attaches the lower limb to the axial skeleton 
through its articulation with the sacrum. The right and 
left hip bones, plus the sacrum and the coccyx, 
together form the pelvis. 


Hip Bone 


The hip bone, or coxal bone, forms the pelvic girdle portion of the pelvis. 
The paired hip bones are the large, curved bones that form the lateral and 
anterior aspects of the pelvis. Each adult hip bone is formed by three 
separate bones that fuse together during the late teenage years. These bony 
components are the ilium, ischium, and pubis ([link]). These names are 
retained and used to define the three regions of the adult hip bone. 

The Hip Bone 


Ilium 


lliac fossa 
lliac crest 
- Posterior 
Posterior Anterior superior 
‘ superior = ‘4 
superior iliac spine iliac spine 
ilac spine a Posterior 
Posterior oe inferior 
inferior weet iliac spine 
iliac spine Acetabulum Auricul rf 
J : uricular surface 
Greater sciatic Arcuate line Greater sciatic 
notch : : 
: Superior ramus of pubis notch 
Ischial body Pubic tubercle Ischial spine 
Ischial spine Lesser sciatic 
Lesser sciatic ont 
notch ; Obturator 
Articular surface foramen 
Ischium of pubis (at pubic Ischium 
symphysis 
Obturator ymphysie) Ischial ramus 
foramen 


Inferior ramus a 
of pubis 


Ischiopubic ramus Ischiopubic ramus 


Ischial tuberosity 
Ischial ramus 


Lateral view, right hip bone Medial view, right hip bone 


The adult hip bone consists of three regions. The ilium 
forms the large, fan-shaped superior portion, the 
ischium forms the posteroinferior portion, and the 
pubis forms the anteromedial portion. 


The ilium is the fan-like, superior region that forms the largest part of the 
hip bone. It is firmly united to the sacrum at the largely immobile sacroiliac 
joint (see [link]). The ischium forms the posteroinferior region of each hip 
bone. It supports the body when sitting. The pubis forms the anterior 
portion of the hip bone. The pubis curves medially, where it joins to the 
pubis of the opposite hip bone at a specialized joint called the pubic 
symphysis. 


Tlium 


When you place your hands on your waist, you can feel the arching, 
superior margin of the ilium along your waistline (see [link]). This curved, 


superior margin of the ilium is the iliac crest. The rounded, anterior 
termination of the iliac crest is the anterior superior iliac spine. This 
important bony landmark can be felt at your anterolateral hip. Inferior to the 
anterior superior iliac spine is a rounded protuberance called the anterior 
inferior iliac spine. Both of these iliac spines serve as attachment points for 
muscles of the thigh. Posteriorly, the iliac crest curves downward to 
terminate as the posterior superior iliac spine. Muscles and ligaments 
surround but do not cover this bony landmark, thus sometimes producing a 
depression seen as a “dimple” located on the lower back. More inferiorly is 
the posterior inferior iliac spine. This is located at the inferior end of a 
large, roughened area called the auricular surface of the ilium. The 
auricular surface articulates with the auricular surface of the sacrum to form 
the sacroiliac joint. Both the posterior superior and posterior inferior iliac 
spines serve as attachment points for the muscles and very strong ligaments 
that support the sacroiliac joint. 


The shallow depression located on the anteromedial (internal) surface of the 
upper ilium is called the iliac fossa. The inferior margin of this space is 
formed by the arcuate line of the ilium, the ridge formed by the 
pronounced change in curvature between the upper and lower portions of 
the ilium. The large, inverted U-shaped indentation located on the posterior 
margin of the lower ilium is called the greater sciatic notch. 


Ischium 


The ischium forms the posterolateral portion of the hip bone (see [link]). 
The large, roughened area of the inferior ischium is the ischial tuberosity. 
This serves as the attachment for the posterior thigh muscles and also 
carries the weight of the body when sitting. You can feel the ischial 
tuberosity if you wiggle your pelvis against the seat of a chair. Projecting 
superiorly and anteriorly from the ischial tuberosity is a narrow segment of 
bone called the ischial ramus. The slightly curved posterior margin of the 
ischium above the ischial tuberosity is the lesser sciatic notch. The bony 
projection separating the lesser sciatic notch and greater sciatic notch is the 
ischial spine. 


Pubis 


The pubis forms the anterior portion of the hip bone (see [link]). The 
enlarged medial portion of the pubis is the pubic body. Located superiorly 
on the pubic body is a small bump called the pubic tubercle. The superior 
pubic ramus is the segment of bone that passes laterally from the pubic 
body to join the ilium. The narrow ridge running along the superior margin 
of the superior pubic ramus is the pectineal line of the pubis. 


The pubic body is joined to the pubic body of the opposite hip bone by the 
pubic symphysis. Extending downward and laterally from the body is the 
inferior pubic ramus. The pubic arch is the bony structure formed by the 
pubic symphysis, and the bodies and inferior pubic rami of the adjacent 
pubic bones. The inferior pubic ramus extends downward to join the ischial 
ramus. Together, these form the single ischiopubic ramus, which extends 
from the pubic body to the ischial tuberosity. The inverted V-shape formed 
as the ischiopubic rami from both sides come together at the pubic 
symphysis is called the subpubic angle. 


Pelvis 


The pelvis consists of four bones: the right and left hip bones, the sacrum, 
and the coccyx (see [link]). The pelvis has several important functions. Its 
primary role is to support the weight of the upper body when sitting and to 
transfer this weight to the lower limbs when standing. It serves as an 
attachment point for trunk and lower limb muscles, and also protects the 
internal pelvic organs. When standing in the anatomical position, the pelvis 
is tilted anteriorly. In this position, the anterior superior iliac spines and the 
pubic tubercles lie in the same vertical plane, and the anterior (internal) 
surface of the sacrum faces forward and downward. 


The three areas of each hip bone, the ilium, pubis, and ischium, converge 
centrally to form a deep, cup-shaped cavity called the acetabulum. This is 
located on the lateral side of the hip bone and is part of the hip joint. The 
large opening in the anteroinferior hip bone between the ischium and pubis 
is the obturator foramen. This space is largely filled in by a layer of 


connective tissue and serves for the attachment of muscles on both its 
internal and external surfaces. 


Several ligaments unite the bones of the pelvis ([link]). The largely 
immobile sacroiliac joint is supported by a pair of strong ligaments that are 
attached between the sacrum and ilium portions of the hip bone. These are 
the anterior sacroiliac ligament on the anterior side of the joint and the 
posterior sacroiliac ligament on the posterior side. Also spanning the 
sacrum and hip bone are two additional ligaments. The sacrospinous 
ligament runs from the sacrum to the ischial spine, and the sacrotuberous 
ligament runs from the sacrum to the ischial tuberosity. These ligaments 
help to support and immobilize the sacrum as it carries the weight of the 
body. 

Ligaments of the Pelvis 


Sacrum 


Posterior 
superior ; 
iliac spine Posterior 

sacroiliac 


" ligament 
llium 9 


Greater 
sciatic 
foramen 


pe rOspInous Sacrospinous 

igament ligament 

Ischial spine 

Pubis Lesser sciatic 
; foramen 

Ischium 

Ischial Sacrotuberous 

tuberosity ligament 


Subpubic 
angle 


Obturator 
foramen 


Ischiopubic ramus 
The posterior sacroiliac ligament supports the 
sacroiliac joint. The sacrospinous ligament spans the 
sacrum to the ischial spine, and the sacrotuberous 
ligament spans the sacrum to the ischial tuberosity. 
The sacrospinous and sacrotuberous ligaments 
contribute to the formation of the greater and lesser 
sciatic foramens. 


Note: 
fepeeeety 
a epenstex COLLEGE 


Watch this video for a 3-D view of the pelvis and its associated ligaments. 
What is the large opening in the bony pelvis, located between the ischium 
and pubic regions, and what two parts of the pubis contribute to the 
formation of this opening? 


The sacrospinous and sacrotuberous ligaments also help to define two 
openings on the posterolateral sides of the pelvis through which muscles, 
nerves, and blood vessels for the lower limb exit. The superior opening is 
the greater sciatic foramen. This large opening is formed by the greater 
sciatic notch of the hip bone, the sacrum, and the sacrospinous ligament. 
The smaller, more inferior lesser sciatic foramen is formed by the lesser 
sciatic notch of the hip bone, together with the sacrospinous and 
sacrotuberous ligaments. 


The space enclosed by the bony pelvis is divided into two regions ({link]). 
The broad, superior region, defined laterally by the large, fan-like portion of 
the upper hip bone, is called the greater pelvis (greater pelvic cavity; false 
pelvis). This broad area is occupied by portions of the small and large 
intestines, and because it is more closely associated with the abdominal 
cavity, it is sometimes referred to as the false pelvis. More inferiorly, the 
narrow, rounded space of the lesser pelvis (lesser pelvic cavity; true pelvis) 
contains the bladder and other pelvic organs, and thus is also known as the 
true pelvis. The pelvic brim (also known as the pelvic inlet) forms the 
superior margin of the lesser pelvis, separating it from the greater pelvis. 
The pelvic brim is defined by a line formed by the upper margin of the 
pubic symphysis anteriorly, and the pectineal line of the pubis, the arcuate 
line of the ilium, and the sacral promontory (the anterior margin of the 


superior sacrum) posteriorly. The inferior limit of the lesser pelvic cavity is 
called the pelvic outlet. This large opening is defined by the inferior margin 
of the pubic symphysis anteriorly, and the ischiopubic ramus, the ischial 
tuberosity, the sacrotuberous ligament, and the inferior tip of the coccyx 
posteriorly. Because of the anterior tilt of the pelvis, the lesser pelvis is also 
angled, giving it an anterosuperior (pelvic inlet) to posteroinferior (pelvic 
outlet) orientation. 

Male and Female Pelvis 


Female 


Subpubic angle 


The female pelvis is adapted for childbirth and is 
broader, with a larger subpubic angle, a rounder pelvic 
brim, and a wider and more shallow lesser pelvic 
cavity than the male pelvis. 


Comparison of the Female and Male Pelvis 


The differences between the adult female and male pelvis relate to function 
and body size. In general, the bones of the male pelvis are thicker and 
heavier, adapted for support of the male’s heavier physical build and 
stronger muscles. The greater sciatic notch of the male hip bone is narrower 
and deeper than the broader notch of females. Because the female pelvis is 
adapted for childbirth, it is wider than the male pelvis, as evidenced by the 
distance between the anterior superior iliac spines (see [link]). The ischial 
tuberosities of females are also farther apart, which increases the size of the 


pelvic outlet. Because of this increased pelvic width, the subpubic angle is 
larger in females (greater than 80 degrees) than it is in males (less than 70 
degrees). The female sacrum is wider, shorter, and less curved, and the 
sacral promontory projects less into the pelvic cavity, thus giving the female 
pelvic inlet (pelvic brim) a more rounded or oval shape compared to males. 
The lesser pelvic cavity of females is also wider and more shallow than the 


narrower, deeper, and tapering lesser pelvis of males. Because of the 

obvious differences between female and male hip bones, this is the one 
bone of the body that allows for the most accurate sex determination. [link] 
provides an overview of the general differences between the female and 


male pelvis. 


Overview of Differences between the Female and Male Pelvis 


Pelvic 
weight 


Pelvic inlet 
shape 


Lesser pelvic 
cavity shape 


Subpubic 
angle 


Pelvic outlet 
shape 


Female pelvis 


Bones of the pelvis 
are lighter and thinner 


Pelvic inlet has a 
round or oval shape 


Lesser pelvic cavity is 
shorter and wider 


Subpubic angle is 
greater than 80 
degrees 


Pelvic outlet is 
rounded and larger 


Male pelvis 


Bones of the pelvis are 
thicker and heavier 


Pelvic inlet is heart- 
shaped 


Lesser pelvic cavity is 
longer and narrower 


Subpubic angle is less 
than 70 degrees 


Pelvic outlet is smaller 


Note: 

Career Connection 

Forensic Pathology and Forensic Anthropology 

A forensic pathologist (also known as a medical examiner) is a medically 
trained physician who has been specifically trained in pathology to 
examine the bodies of the deceased to determine the cause of death. A 
forensic pathologist applies his or her understanding of disease as well as 
toxins, blood and DNA analysis, firearms and ballistics, and other factors 
to assess the cause and manner of death. At times, a forensic pathologist 
will be called to testify under oath in situations that involve a possible 
crime. Forensic pathology is a field that has received much media attention 
on television shows or following a high-profile death. 

While forensic pathologists are responsible for determining whether the 
cause of someone’s death was natural, a suicide, accidental, or a homicide, 
there are times when uncovering the cause of death is more complex, and 
other skills are needed. Forensic anthropology brings the tools and 
knowledge of physical anthropology and human osteology (the study of the 
skeleton) to the task of investigating a death. A forensic anthropologist 
assists medical and legal professionals in identifying human remains. The 
science behind forensic anthropology involves the study of archaeological 
excavation; the examination of hair; an understanding of plants, insects, 
and footprints; the ability to determine how much time has elapsed since 
the person died; the analysis of past medical history and toxicology; the 
ability to determine whether there are any postmortem injuries or 
alterations of the skeleton; and the identification of the decedent (deceased 
person) using skeletal and dental evidence. 

Due to the extensive knowledge and understanding of excavation 
techniques, a forensic anthropologist is an integral and invaluable team 
member to have on-site when investigating a crime scene, especially when 
the recovery of human skeletal remains is involved. When remains are 
bought to a forensic anthropologist for examination, he or she must first 
determine whether the remains are in fact human. Once the remains have 
been identified as belonging to a person and not to an animal, the next step 
is to approximate the individual’s age, sex, race, and height. The forensic 
anthropologist does not determine the cause of death, but rather provides 
information to the forensic pathologist, who will use all of the data 
collected to make a final determination regarding the cause of death. 


Chapter Review 


The pelvic girdle, consisting of a hip bone, serves to attach a lower limb to 
the axial skeleton. The hip bone articulates posteriorly at the sacroiliac joint 
with the sacrum, which is part of the axial skeleton. The right and left hip 
bones converge anteriorly and articulate with each other at the pubic 
symphysis. The combination of the hip bone, the sacrum, and the coccyx 
forms the pelvis. The pelvis has a pronounced anterior tilt. The primary 
function of the pelvis is to support the upper body and transfer body weight 
to the lower limbs. It also serves as the site of attachment for multiple 
muscles. 


The hip bone consists of three regions: the ilium, ischium, and pubis. The 
ilium forms the large, fan-like region of the hip bone. The superior margin 
of this area is the iliac crest. Located at either end of the iliac crest are the 
anterior superior and posterior superior iliac spines. Inferior to these are the 
anterior inferior and posterior inferior iliac spines. The auricular surface of 
the ilium articulates with the sacrum to form the sacroiliac joint. The medial 
surface of the upper ilium forms the iliac fossa, with the arcuate line 
marking the inferior limit of this area. The posterior margin of the ilium has 
the large greater sciatic notch. 


The posterolateral portion of the hip bone is the ischium. It has the 
expanded ischial tuberosity, which supports body weight when sitting. The 
ischial ramus projects anteriorly and superiorly. The posterior margin of the 
ischium has the shallow lesser sciatic notch and the ischial spine, which 
separates the greater and lesser sciatic notches. 


The pubis forms the anterior portion of the hip bone. The body of the pubis 
articulates with the pubis of the opposite hip bone at the pubic symphysis. 
The superior margin of the pubic body has the pubic tubercle. The pubis is 
joined to the ilium by the superior pubic ramus, the superior surface of 
which forms the pectineal line. The inferior pubic ramus projects inferiorly 
and laterally. The pubic arch is formed by the pubic symphysis, the bodies 
of the adjacent pubic bones, and the two inferior pubic rami. The inferior 
pubic ramus joins the ischial ramus to form the ischiopubic ramus. The 


subpubic angle is formed by the medial convergence of the right and left 
ischiopubic rami. 


The lateral side of the hip bone has the cup-like acetabulum, which is part 
of the hip joint. The large anterior opening is the obturator foramen. The 
sacroiliac joint is supported by the anterior and posterior sacroiliac 
ligaments. The sacrum is also joined to the hip bone by the sacrospinous 
ligament, which attaches to the ischial spine, and the sacrotuberous 
ligament, which attaches to the ischial tuberosity. The sacrospinous and 
sacrotuberous ligaments contribute to the formation of the greater and lesser 
sciatic foramina. 


The broad space of the upper pelvis is the greater pelvis, and the narrow, 
inferior space is the lesser pelvis. These areas are separated by the pelvic 
brim (pelvic inlet). The inferior opening of the pelvis is the pelvic outlet. 
Compared to the male, the female pelvis is wider to accommodate 
childbirth, has a larger subpubic angle, and a broader greater sciatic notch. 


Interactive Link Questions 


Exercise: 
Problem: 
Watch this video for a 3-D view of the pelvis and its associated 
ligaments. What is the large opening in the bony pelvis, located 


between the ischium and pubic regions, and what two parts of the 
pubis contribute to the formation of this opening? 


Solution: 


The obturator foramen is located between the ischium and the pubis. 
The superior and inferior pubic rami contribute to the boundaries of 
the obturator foramen. 


Review Questions 


Exercise: 


Problem:How many bones fuse in adulthood to form the hip bone? 


Boop 
ul B WN 


Solution: 


B 


Exercise: 


Problem: Which component forms the superior part of the hip bone? 


a. ilium 
b. pubis 
c. ischium 
d. sacrum 


Solution: 


A 


Exercise: 


Problem: Which of the following supports body weight when sitting? 


a. iliac crest 

b. ischial tuberosity 
c. ischiopubic ramus 
d. pubic body 


Solution: 


B 
Exercise: 


Problem: 
The ischial spine is found between which of the following structures? 


a. inferior pubic ramus and ischial ramus 

b. pectineal line and arcuate line 

c. lesser sciatic notch and greater sciatic notch 

d. anterior superior iliac spine and posterior superior iliac spine 


Solution: 


C 


Exercise: 


Problem:The pelvis 


a. has a subpubic angle that is larger in females 

b. consists of the two hip bones, but does not include the sacrum or 
coccyx 

c. has an obturator foramen, an opening that is defined in part by the 
Sacrospinous and sacrotuberous ligaments 

d. has a space located inferior to the pelvic brim called the greater 
pelvis 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: 


Describe the articulations and ligaments that unite the four bones of 
the pelvis to each other. 


Solution: 


The pelvis is formed by the combination of the right and left hip 
bones, the sacrum, and the coccyx. The auricular surfaces of each hip 
bone articulate with the auricular surface of the sacrum to form the 
sacroiliac joint. This joint is supported on either side by the strong 
anterior and posterior sacroiliac ligaments. The right and left hip bones 
converge anteriorly, where the pubic bodies articulate with each other 
to form the pubic symphysis joint. The sacrum is also attached to the 
hip bone by the sacrospinous ligament, which spans the sacrum to the 
ischial spine, and the sacrotuberous ligament, which runs from the 
sacrum to the ischial tuberosity. The coccyx is attached to the inferior 
end of the sacrum. 


Exercise: 


Problem: 
Discuss the ways in which the female pelvis is adapted for childbirth. 
Solution: 


Compared to the male, the female pelvis is wider to accommodate 
childbirth. Thus, the female pelvis has greater distances between the 
anterior superior iliac spines and between the ischial tuberosities. The 
greater width of the female pelvis results in a larger subpubic angle. 
This angle, formed by the anterior convergence of the right and left 
ischiopubic rami, is larger in females (greater than 80 degrees) than in 
males (less than 70 degrees). The female sacral promontory does not 
project anteriorly as far as it does in males, which gives the pelvic 
brim (pelvic inlet) of the female a rounded or oval shape. The lesser 
pelvic cavity is wider and more shallow in females, and the pelvic 


outlet is larger than in males. Thus, the greater width of the female 
pelvis, with its larger pelvic inlet, lesser pelvis, and pelvic outlet, are 
important for childbirth because the baby must pass through the pelvis 
during delivery. 


Glossary 


acetabulum 
large, cup-shaped cavity located on the lateral side of the hip bone; 
formed by the junction of the ilium, pubis, and ischium portions of the 
hip bone 


anterior inferior iliac spine 
small, bony projection located on the anterior margin of the ilium, 
below the anterior superior iliac spine 


anterior sacroiliac ligament 
strong ligament between the sacrum and the ilium portions of the hip 
bone that supports the anterior side of the sacroiliac joint 


anterior superior iliac spine 
rounded, anterior end of the iliac crest 


arcuate line of the ilium 
smooth ridge located at the inferior margin of the iliac fossa; forms the 
lateral portion of the pelvic brim 


auricular surface of the ilium 
roughened area located on the posterior, medial side of the ilium of the 
hip bone; articulates with the auricular surface of the sacrum to form 
the sacroiliac joint 


coxal bone 
hip bone 


greater pelvis 


(also, greater pelvic cavity or false pelvis) broad space above the 
pelvic brim defined laterally by the fan-like portion of the upper ilium 


greater sciatic foramen 
pelvic opening formed by the greater sciatic notch of the hip bone, the 
sacrum, and the sacrospinous ligament 


greater Sciatic notch 
large, U-shaped indentation located on the posterior margin of the 
ilium, superior to the ischial spine 


hip bone 
coxal bone; single bone that forms the pelvic girdle; consists of three 
areas, the ilium, ischium, and pubis 


iliac crest 
curved, superior margin of the ilium 


iliac fossa 
shallow depression found on the anterior and medial surfaces of the 
upper ilium 


ilium 
superior portion of the hip bone 


inferior pubic ramus 
narrow segment of bone that passes inferiorly and laterally from the 
pubic body; joins with the ischial ramus to form the ischiopubic ramus 


ischial ramus 
bony extension projecting anteriorly and superiorly from the ischial 
tuberosity; joins with the inferior pubic ramus to form the ischiopubic 
ramus 


ischial spine 
pointed, bony projection from the posterior margin of the ischium that 
separates the greater sciatic notch and lesser sciatic notch 


ischial tuberosity 
large, roughened protuberance that forms the posteroinferior portion of 
the hip bone; weight-bearing region of the pelvis when sitting 


ischiopubic ramus 
narrow extension of bone that connects the ischial tuberosity to the 
pubic body; formed by the junction of the ischial ramus and inferior 
pubic ramus 


ischium 
posteroinferior portion of the hip bone 


lesser pelvis 
(also, lesser pelvic cavity or true pelvis) narrow space located within 
the pelvis, defined superiorly by the pelvic brim (pelvic inlet) and 
inferiorly by the pelvic outlet 


lesser sciatic foramen 
pelvic opening formed by the lesser sciatic notch of the hip bone, the 
Sacrospinous ligament, and the sacrotuberous ligament 


lesser sciatic notch 
shallow indentation along the posterior margin of the ischium, inferior 
to the ischial spine 


obturator foramen 
large opening located in the anterior hip bone, between the pubis and 
ischium regions 


pectineal line 
narrow ridge located on the superior surface of the superior pubic 
ramus 


pelvic brim 
pelvic inlet; the dividing line between the greater and lesser pelvic 
regions; formed by the superior margin of the pubic symphysis, the 
pectineal lines of each pubis, the arcuate lines of each ilium, and the 
sacral promontory 


pelvic girdle 
hip girdle; consists of a single hip bone, which attaches a lower limb to 
the sacrum of the axial skeleton 


pelvic inlet 
pelvic brim 


pelvic outlet 
inferior opening of the lesser pelvis; formed by the inferior margin of 
the pubic symphysis, right and left ischiopubic rami and sacrotuberous 
ligaments, and the tip of the coccyx 


pelvis 
ring of bone consisting of the right and left hip bones, the sacrum, and 
the coccyx 


posterior inferior iliac spine 
small, bony projection located at the inferior margin of the auricular 
surface on the posterior ilium 


posterior sacroiliac ligament 
strong ligament spanning the sacrum and ilium of the hip bone that 
supports the posterior side of the sacroiliac joint 


posterior superior iliac spine 
rounded, posterior end of the iliac crest 


pubic arch 
bony structure formed by the pubic symphysis, and the bodies and 
inferior pubic rami of the right and left pubic bones 


pubic body 
enlarged, medial portion of the pubis region of the hip bone 


pubic symphysis 
joint formed by the articulation between the pubic bodies of the right 
and left hip bones 


pubic tubercle 
small bump located on the superior aspect of the pubic body 


pubis 
anterior portion of the hip bone 


sacroiliac joint 
joint formed by the articulation between the auricular surfaces of the 
sacrum and ilium 


sacrospinous ligament 
ligament that spans the sacrum to the ischial spine of the hip bone 


sacrotuberous ligament 
ligament that spans the sacrum to the ischial tuberosity of the hip bone 


subpubic angle 
inverted V-shape formed by the convergence of the right and left 
ischiopubic rami; this angle is greater than 80 degrees in females and 
less than 70 degrees in males 


superior pubic ramus 
narrow segment of bone that passes laterally from the pubic body to 
join the ilium 


Bones of the Lower Limb 
By the end of this section, you will be able to: 


e Identify the divisions of the lower limb and describe the bones of each 
region 

e Describe the bones and bony landmarks that articulate at each joint of 
the lower limb 


Like the upper limb, the lower limb is divided into three regions. The thigh 
is that portion of the lower limb located between the hip joint and knee 
joint. The leg is specifically the region between the knee joint and the ankle 
joint. Distal to the ankle is the foot. The lower limb contains 30 bones. 
These are the femur, patella, tibia, fibula, tarsal bones, metatarsal bones, 
and phalanges (see [link]). The femur is the single bone of the thigh. The 
patella is the kneecap and articulates with the distal femur. The tibia is the 
larger, weight-bearing bone located on the medial side of the leg, and the 
fibula is the thin bone of the lateral leg. The bones of the foot are divided 
into three groups. The posterior portion of the foot is formed by a group of 
seven bones, each of which is known as a tarsal bone, whereas the mid- 
foot contains five elongated bones, each of which is a metatarsal bone. 
The toes contain 14 small bones, each of which is a phalanx bone of the 
foot. 


Femur 


The femur, or thigh bone, is the single bone of the thigh region ((link]). It is 
the longest and strongest bone of the body, and accounts for approximately 
one-quarter of a person’s total height. The rounded, proximal end is the 
head of the femur, which articulates with the acetabulum of the hip bone 
to form the hip joint. The fovea capitis is a minor indentation on the 
medial side of the femoral head that serves as the site of attachment for the 
ligament of the head of the femur. This ligament spans the femur and 
acetabulum, but is weak and provides little support for the hip joint. It does, 
however, carry an important artery that supplies the head of the femur. 
Femur and Patella 


Greater 
trochanter 


Greater 
trochanter 


Gluteal 
tuberosity 


Intertrochanteric line 


Intertrochanteric 
crest 


Lesser trochanter 
Linea 
aspera 


Adductor 


tubercle Lateral 


Lateral epicondyle 


epicondyle Medial 
oe Intercondylar 
Patella fossa 
Medial condyle Lateral 
i condyle 
Fibula — 
Fibula 


Anterior view Posterior view 


The femur is the single bone of the thigh 
region. It articulates superiorly with the 
hip bone at the hip joint, and inferiorly 

with the tibia at the knee joint. The patella 
only articulates with the distal end of the 
femur. 


The narrowed region below the head is the neck of the femur. This is a 
common area for fractures of the femur. The greater trochanter is the 
large, upward, bony projection located above the base of the neck. Multiple 
muscles that act across the hip joint attach to the greater trochanter, which, 
because of its projection from the femur, gives additional leverage to these 
muscles. The greater trochanter can be felt just under the skin on the lateral 
side of your upper thigh. The lesser trochanter is a small, bony 
prominence that lies on the medial aspect of the femur, just below the neck. 
A single, powerful muscle attaches to the lesser trochanter. Running 
between the greater and lesser trochanters on the anterior side of the femur 
is the roughened intertrochanteric line. The trochanters are also connected 
on the posterior side of the femur by the larger intertrochanteric crest. 


The elongated shaft of the femur has a slight anterior bowing or curvature. 
At its proximal end, the posterior shaft has the gluteal tuberosity, a 
roughened area extending inferiorly from the greater trochanter. More 
inferiorly, the gluteal tuberosity becomes continuous with the linea aspera 
(“rough line”). This is the roughened ridge that passes distally along the 
posterior side of the mid-femur. Multiple muscles of the hip and thigh 
regions make long, thin attachments to the femur along the linea aspera. 


The distal end of the femur has medial and lateral bony expansions. On the 
lateral side, the smooth portion that covers the distal and posterior aspects 
of the lateral expansion is the lateral condyle of the femur. The roughened 
area on the outer, lateral side of the condyle is the lateral epicondyle of the 
femur. Similarly, the smooth region of the distal and posterior medial 
femur is the medial condyle of the femur, and the irregular outer, medial 
side of this is the medial epicondyle of the femur. The lateral and medial 
condyles articulate with the tibia to form the knee joint. The epicondyles 
provide attachment for muscles and supporting ligaments of the knee. The 
adductor tubercle is a small bump located at the superior margin of the 
medial epicondyle. Posteriorly, the medial and lateral condyles are 
separated by a deep depression called the intercondylar fossa. Anteriorly, 
the smooth surfaces of the condyles join together to form a wide groove 
called the patellar surface, which provides for articulation with the patella 
bone. The combination of the medial and lateral condyles with the patellar 
surface gives the distal end of the femur a horseshoe (U) shape. 


Watch this video to view how a fracture of the mid-femur is surgically 
repaired. How are the two portions of the broken femur stabilized during 
surgical repair of a fractured femur? 


Patella 


The patella (kneecap) is largest sesamoid bone of the body (see [link]). A 
sesamoid bone is a bone that is incorporated into the tendon of a muscle 
where that tendon crosses a joint. The sesamoid bone articulates with the 
underlying bones to prevent damage to the muscle tendon due to rubbing 
against the bones during movements of the joint. The patella is found in the 
tendon of the quadriceps femoris muscle, the large muscle of the anterior 
thigh that passes across the anterior knee to attach to the tibia. The patella 
articulates with the patellar surface of the femur and thus prevents rubbing 
of the muscle tendon against the distal femur. The patella also lifts the 
tendon away from the knee joint, which increases the leverage power of the 
quadriceps femoris muscle as it acts across the knee. The patella does not 
articulate with the tibia. 


Note: 


openstax COLLEGE 
. a 


| r 


Pit | | 


Visit this site to perform a virtual knee replacement surgery. The prosthetic 
knee components must be properly aligned to function properly. How is 
this alignment ensured? 


Note: 

Homeostatic Imbalances 

Runner’s Knee 

Runner’s knee, also known as patellofemoral syndrome, is the most 
common overuse injury among runners. It is most frequent in adolescents 
and young adults, and is more common in females. It often results from 
excessive running, particularly downhill, but may also occur in athletes 
who do a lot of knee bending, such as jumpers, skiers, cyclists, weight 
lifters, and soccer players. It is felt as a dull, aching pain around the front 
of the knee and deep to the patella. The pain may be felt when walking or 
running, going up or down stairs, kneeling or squatting, or after sitting with 
the knee bent for an extended period. 

Patellofemoral syndrome may be initiated by a variety of causes, including 
individual variations in the shape and movement of the patella, a direct 
blow to the patella, or flat feet or improper shoes that cause excessive 
turning in or out of the feet or leg. These factors may cause in an 
imbalance in the muscle pull that acts on the patella, resulting in an 
abnormal tracking of the patella that allows it to deviate too far toward the 
lateral side of the patellar surface on the distal femur. 

Because the hips are wider than the knee region, the femur has a diagonal 
orientation within the thigh, in contrast to the vertically oriented tibia of 
the leg ({link]). The Q-angle is a measure of how far the femur is angled 
laterally away from vertical. The Q-angle is normally 10—15 degrees, with 
females typically having a larger Q-angle due to their wider pelvis. During 
extension of the knee, the quadriceps femoris muscle pulls the patella both 


superiorly and laterally, with the lateral pull greater in women due to their 
large Q-angle. This makes women more vulnerable to developing 
patellofemoral syndrome than men. Normally, the large lip on the lateral 
side of the patellar surface of the femur compensates for the lateral pull on 
the patella, and thus helps to maintain its proper tracking. 

However, if the pull produced by the medial and lateral sides of the 
quadriceps femoris muscle is not properly balanced, abnormal tracking of 
the patella toward the lateral side may occur. With continued use, this 
produces pain and could result in damage to the articulating surfaces of the 
patella and femur, and the possible future development of arthritis. 
Treatment generally involves stopping the activity that produces knee pain 
for a period of time, followed by a gradual resumption of activity. Proper 
strengthening of the quadriceps femoris muscle to correct for imbalances is 
also important to help prevent reoccurrence. 

The Q-Angle 


Q-angle 


Anterior view 


The Q-angle is a 
measure of the amount 
of lateral deviation of 
the femur from the 
vertical line of the 
tibia. Adult females 
have a larger Q-angle 
due to their wider 
pelvis than adult 
males. 


Tibia 


The tibia (shin bone) is the medial bone of the leg and is larger than the 
fibula, with which it is paired ({link]). The tibia is the main weight-bearing 
bone of the lower leg and the second longest bone of the body, after the 
femur. The medial side of the tibia is located immediately under the skin, 
allowing it to be easily palpated down the entire length of the medial leg. 
Tibia and Fibula 


fa a 


Lateral condyle : 
Articular surface 


of lateral condyle 


Articular surface 
of medial condyle 


Medial Head of fibula 


aa ia Anterior border 


Interosseous 
membrane 


Tibial tuberosity 


Soleal line 


Fibula Fibula 


Medial malleolus 


Lateral malleolus 
Lateral malleolus 


Articular surface 
Anterior view Posterior view 


The tibia is the larger, weight-bearing bone 
located on the medial side of the leg. The 
fibula is the slender bone of the lateral side 
of the leg and does not bear weight. 


The proximal end of the tibia is greatly expanded. The two sides of this 
expansion form the medial condyle of the tibia and the lateral condyle of 
the tibia. The tibia does not have epicondyles. The top surface of each 
condyle is smooth and flattened. These areas articulate with the medial and 
lateral condyles of the femur to form the knee joint. Between the 
articulating surfaces of the tibial condyles is the intercondylar eminence, 
an irregular, elevated area that serves as the inferior attachment point for 
two supporting ligaments of the knee. 


The tibial tuberosity is an elevated area on the anterior side of the tibia, 
near its proximal end. It is the final site of attachment for the muscle tendon 
associated with the patella. More inferiorly, the shaft of the tibia becomes 
triangular in shape. The anterior apex of 


MH this triangle forms the anterior border of the tibia, which begins at 
the tibial tuberosity and runs inferiorly along the length of the tibia. Both 
the anterior border and the medial side of the triangular shaft are located 
immediately under the skin and can be easily palpated along the entire 
length of the tibia. A small ridge running down the lateral side of the tibial 
shaft is the interosseous border of the tibia. This is for the attachment of 
the interosseous membrane of the leg, the sheet of dense connective tissue 
that unites the tibia and fibula bones. Located on the posterior side of the 
tibia is the soleal line, a diagonally running, roughened ridge that begins 
below the base of the lateral condyle, and runs down and medially across 
the proximal third of the posterior tibia. Muscles of the posterior leg attach 
to this line. 


The large expansion found on the medial side of the distal tibia is the 
medial malleolus (“little hammer”). This forms the large bony bump found 
on the medial side of the ankle region. Both the smooth surface on the 
inside of the medial malleolus and the smooth area at the distal end of the 
tibia articulate with the talus bone of the foot as part of the ankle joint. On 
the lateral side of the distal tibia is a wide groove called the fibular notch. 
This area articulates with the distal end of the fibula, forming the distal 
tibiofibular joint. 


Fibula 


The fibula is the slender bone located on the lateral side of the leg (see 
[link]). The fibula does not bear weight. It serves primarily for muscle 
attachments and thus is largely surrounded by muscles. Only the proximal 
and distal ends of the fibula can be palpated. 


The head of the fibula is the small, knob-like, proximal end of the fibula. It 
articulates with the inferior aspect of the lateral tibial condyle, forming the 
proximal tibiofibular joint. The thin shaft of the fibula has the 
interosseous border of the fibula, a narrow ridge running down its medial 
side for the attachment of the interosseous membrane that spans the fibula 
and tibia. The distal end of the fibula forms the lateral malleolus, which 
forms the easily palpated bony bump on the lateral side of the ankle. The 
deep (medial) side of the lateral malleolus articulates with the talus bone of 
the foot as part of the ankle joint. The distal fibula also articulates with the 
fibular notch of the tibia. 


Tarsal Bones 


The posterior half of the foot is formed by seven tarsal bones ([link]). The 
most superior bone is the talus. This has a relatively square-shaped, upper 
surface that articulates with the tibia and fibula to form the ankle joint. 
Three areas of articulation form the ankle joint: The superomedial surface 
of the talus bone articulates with the medial malleolus of the tibia, the top of 
the talus articulates with the distal end of the tibia, and the lateral side of the 
talus articulates with the lateral malleolus of the fibula. Inferiorly, the talus 
articulates with the calcaneus (heel bone), the largest bone of the foot, 
which forms the heel. Body weight is transferred from the tibia to the talus 
to the calcaneus, which rests on the ground. The medial calcaneus has a 
prominent bony extension called the sustentaculum tali (“support for the 
talus”) that supports the medial side of the talus bone. 

Bones of the Foot 


| \ t | Navicular Talus 
\ Intermediate 
Tarsals cuneiform 


Metatarsals 
Phalanges First metatarsal 


Facet for medial 
malleolus 


Sustentaculum 
tali (talar shelf) 


Distal ————___{ : 
¢€ Medial Calcaneus 

Proximal Y Distal cuneiform =i ee 
halanges phalange alcaneal tuberosi 
i 9 : / Medial view 

Middle 

phalange 
Medial Proximal 
cuneiform phalange Facet for 

lateral malleolus Navicular Intermediate cuneiform 


Intermediate 
cuneiform 


Lateral 


cuneiform Lateral cuneiform 


Navicular Cuboid 


Talus 


Trochlea 


of talus 
Calcaneus 


Calcaneus Cuboid Fifth metatarsal 


Superior view Lateral view 


The bones of the foot are divided into three groups. The posterior 
foot is formed by the seven tarsal bones. The mid-foot has the five 
metatarsal bones. The toes contain the phalanges. 


The cuboid bone articulates with the anterior end of the calcaneus bone. 
The cuboid has a deep groove running across its inferior surface, which 
provides passage for a muscle tendon. The talus bone articulates anteriorly 
with the navicular bone, which in turn articulates anteriorly with the three 
cuneiform (“wedge-shaped”) bones. These bones are the medial 
cuneiform, the intermediate cuneiform, and the lateral cuneiform. Each 
of these bones has a broad superior surface and a narrow inferior surface, 
which together produce the transverse (medial-lateral) curvature of the foot. 
The navicular and lateral cuneiform bones also articulate with the medial 
side of the cuboid bone. 


Note: 
— openstax COLLEGE 
1 


Oe 


Use this tutorial to review the bones of the foot. Which tarsal bones are in 
the proximal, intermediate, and distal groups? 


Metatarsal Bones 


The anterior half of the foot is formed by the five metatarsal bones, which 
are located between the tarsal bones of the posterior foot and the phalanges 
of the toes (see [link]). These elongated bones are numbered 1-5, starting 
with the medial side of the foot. The first metatarsal bone is shorter and 
thicker than the others. The second metatarsal is the longest. The base of 
the metatarsal bone is the proximal end of each metatarsal bone. These 
articulate with the cuboid or cuneiform bones. The base of the fifth 
metatarsal has a large, lateral expansion that provides for muscle 
attachments. This expanded base of the fifth metatarsal can be felt as a bony 
bump at the midpoint along the lateral border of the foot. The expanded 
distal end of each metatarsal is the head of the metatarsal bone. Each 
metatarsal bone articulates with the proximal phalanx of a toe to forma 
metatarsophalangeal joint. The heads of the metatarsal bones also rest on 
the ground and form the ball (anterior end) of the foot. 


Phalanges 


The toes contain a total of 14 phalanx bones (phalanges), arranged in a 
similar manner as the phalanges of the fingers (see [link]). The toes are 
numbered 1-5, starting with the big toe (hallux). The big toe has two 
phalanx bones, the proximal and distal phalanges. The remaining toes all 


have proximal, middle, and distal phalanges. A joint between adjacent 
phalanx bones is called an interphalangeal joint. 


Note: 


eS 


wees Openstax COLLEGE 


aha 


View this link to learn about a bunion, a localized swelling on the medial 
side of the foot, next to the first metatarsophalangeal joint, at the base of 
the big toe. What is a bunion and what type of shoe is most likely to cause 
this to develop? 


Arches of the Foot 


When the foot comes into contact with the ground during walking, running, 
or jumping activities, the impact of the body weight puts a tremendous 
amount of pressure and force on the foot. During running, the force applied 
to each foot as it contacts the ground can be up to 2.5 times your body 
weight. The bones, joints, ligaments, and muscles of the foot absorb this 
force, thus greatly reducing the amount of shock that is passed superiorly 
into the lower limb and body. The arches of the foot play an important role 
in this shock-absorbing ability. When weight is applied to the foot, these 
arches will flatten somewhat, thus absorbing energy. When the weight is 
removed, the arch rebounds, giving “spring” to the step. The arches also 
serve to distribute body weight side to side and to either end of the foot. 


The foot has a transverse arch, a medial longitudinal arch, and a lateral 
longitudinal arch (see [link]). The transverse arch forms the medial-lateral 
curvature of the mid-foot. It is formed by the wedge shapes of the 
cuneiform bones and bases (proximal ends) of the first to fourth metatarsal 


bones. This arch helps to distribute body weight from side to side within the 
foot, thus allowing the foot to accommodate uneven terrain. 


The longitudinal arches run down the length of the foot. The lateral 
longitudinal arch is relatively flat, whereas the medial longitudinal arch is 
larger (taller). The longitudinal arches are formed by the tarsal bones 
posteriorly and the metatarsal bones anteriorly. These arches are supported 
at either end, where they contact the ground. Posteriorly, this support is 
provided by the calcaneus bone and anteriorly by the heads (distal ends) of 
the metatarsal bones. The talus bone, which receives the weight of the body, 
is located at the top of the longitudinal arches. Body weight is then 
conveyed from the talus to the ground by the anterior and posterior ends of 
these arches. Strong ligaments unite the adjacent foot bones to prevent 
disruption of the arches during weight bearing. On the bottom of the foot, 
additional ligaments tie together the anterior and posterior ends of the 
arches. These ligaments have elasticity, which allows them to stretch 
somewhat during weight bearing, thus allowing the longitudinal arches to 
spread. The stretching of these ligaments stores energy within the foot, 
rather than passing these forces into the leg. Contraction of the foot muscles 
also plays an important role in this energy absorption. When the weight is 
removed, the elastic ligaments recoil and pull the ends of the arches closer 
together. This recovery of the arches releases the stored energy and 
improves the energy efficiency of walking. 


Stretching of the ligaments that support the longitudinal arches can lead to 
pain. This can occur in overweight individuals, with people who have jobs 
that involve standing for long periods of time (such as a waitress), or 
walking or running long distances. If stretching of the ligaments is 
prolonged, excessive, or repeated, it can result in a gradual lengthening of 
the supporting ligaments, with subsequent depression or collapse of the 
longitudinal arches, particularly on the medial side of the foot. This 
condition is called pes planus (“flat foot” or “fallen arches”). 


Chapter Review 


The lower limb is divided into three regions. These are the thigh, located 
between the hip and knee joints; the leg, located between the knee and 


ankle joints; and distal to the ankle, the foot. There are 30 bones in each 
lower limb. These are the femur, patella, tibia, fibula, seven tarsal bones, 
five metatarsal bones, and 14 phalanges. 


The femur is the single bone of the thigh. Its rounded head articulates with 
the acetabulum of the hip bone to form the hip joint. The head has the fovea 
capitis for attachment of the ligament of the head of the femur. The narrow 
neck joins inferiorly with the greater and lesser trochanters. Passing 
between these bony expansions are the intertrochanteric line on the anterior 
femur and the larger intertrochanteric crest on the posterior femur. On the 
posterior shaft of the femur is the gluteal tuberosity proximally and the 
linea aspera in the mid-shaft region. The expanded distal end consists of 
three articulating surfaces: the medial and lateral condyles, and the patellar 
surface. The outside margins of the condyles are the medial and lateral 
epicondyles. The adductor tubercle is on the superior aspect of the medial 
epicondyle. 


The patella is a sesamoid bone located within a muscle tendon. It articulates 
with the patellar surface on the anterior side of the distal femur, thereby 
protecting the muscle tendon from rubbing against the femur. 


The leg contains the large tibia on the medial side and the slender fibula on 
the lateral side. The tibia bears the weight of the body, whereas the fibula 
does not bear weight. The interosseous border of each bone is the 
attachment site for the interosseous membrane of the leg, the connective 
tissue sheet that unites the tibia and fibula. 


The proximal tibia consists of the expanded medial and lateral condyles, 
which articulate with the medial and lateral condyles of the femur to form 
the knee joint. Between the tibial condyles is the intercondylar eminence. 
On the anterior side of the proximal tibia is the tibial tuberosity, which is 
continuous inferiorly with the anterior border of the tibia. On the posterior 
side, the proximal tibia has the curved soleal line. The bony expansion on 
the medial side of the distal tibia is the medial malleolus. The groove on the 
lateral side of the distal tibia is the fibular notch. 


The head of the fibula forms the proximal end and articulates with the 
underside of the lateral condyle of the tibia. The distal fibula articulates 


with the fibular notch of the tibia. The expanded distal end of the fibula is 
the lateral malleolus. 


The posterior foot is formed by the seven tarsal bones. The talus articulates 
superiorly with the distal tibia, the medial malleolus of the tibia, and the 
lateral malleolus of the fibula to form the ankle joint. The talus articulates 
inferiorly with the calcaneus bone. The sustentaculum tali of the calcaneus 
helps to support the talus. Anterior to the talus is the navicular bone, and 
anterior to this are the medial, intermediate, and lateral cuneiform bones. 
The cuboid bone is anterior to the calcaneus. 


The five metatarsal bones form the anterior foot. The base of these bones 
articulate with the cuboid or cuneiform bones. The metatarsal heads, at their 
distal ends, articulate with the proximal phalanges of the toes. The big toe 
(toe number 1) has proximal and distal phalanx bones. The remaining toes 
have proximal, middle, and distal phalanges. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to view how a fracture of the mid-femur is surgically 
repaired. How are the two portions of the broken femur stabilized 
during surgical repair of a fractured femur? 


Solution: 


A hole is drilled into the greater trochanter, the bone marrow 
(medullary) space inside the femur is enlarged, and finally an 
intramedullary rod is inserted into the femur. This rod is then anchored 
to the bone with screws. 


Exercise: 


Problem: 


Visit this site to perform a virtual knee replacement surgery. The 
prosthetic knee components must be properly aligned to function 
properly. How is this alignment ensured? 


Solution: 

Metal cutting jigs are attached to the bones to ensure that the bones are 

cut properly prior to the attachment of prosthetic components. 
Exercise: 

Problem: 


Use this tutorial to review the bones of the foot. Which tarsal bones are 
in the proximal, intermediate, and distal groups? 


Solution: 


The proximal group of tarsal bones includes the calcaneus and talus 
bones, the navicular bone is intermediate, and the distal group consists 
of the cuboid bone plus the medial, intermediate, and lateral cuneiform 
bones. 


Exercise: 
Problem: 
View this link to learn about a bunion, a localized swelling on the 
medial side of the foot, next to the first metatarsophalangeal joint, at 


the base of the big toe. What is a bunion and what type of shoe is most 
likely to cause this to develop? 


Solution: 
A bunion results from the deviation of the big toe toward the second 


toe, which causes the distal end of the first metatarsal bone to stick out. 
A bunion may also be caused by prolonged pressure on the foot from 


pointed shoes with a narrow toe box that compresses the big toe and 
pushes it toward the second toe. 


Review Questions 


Exercise: 


Problem: 


Which bony landmark of the femur serves as a site for muscle 
attachments? 


a. fovea capitis 

b. lesser trochanter 
c. head 

d. medial condyle 


Solution: 


B 


Exercise: 


Problem: What structure contributes to the knee joint? 


a. lateral malleolus of the fibula 
b. tibial tuberosity 

c. medial condyle of the tibia 

d. lateral epicondyle of the femur 


Solution: 


C 


Exercise: 


Problem: Which tarsal bone articulates with the tibia and fibula? 


a. calcaneus 
b. cuboid 

c. navicular 
d. talus 


Solution: 


D 


Exercise: 


Problem: What is the total number of bones found in the foot and toes? 


a. 7 


a 
aN 
BS 


Solution: 


C 


Exercise: 


Problem: The tibia 


a. has an expanded distal end called the lateral malleolus 

b. is not a weight-bearing bone 

c. is firmly anchored to the fibula by an interosseous membrane 

d. can be palpated (felt) under the skin only at its proximal and 
distal ends 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: 


Define the regions of the lower limb, name the bones found in each 
region, and describe the bony landmarks that articulate together to 
form the hip, knee, and ankle joints. 


Solution: 


The lower limb is divided into three regions. The thigh is the region 
located between the hip and knee joints. It contains the femur and the 
patella. The hip joint is formed by the articulation between the 
acetabulum of the hip bone and the head of the femur. The leg is the 
region between the knee and ankle joints, and contains the tibia 
(medially) and the fibula (laterally). The knee joint is formed by the 
articulations between the medial and lateral condyles of the femur, and 
the medial and lateral condyles of the tibia. Also associated with the 
knee is the patella, which articulates with the patellar surface of the 
distal femur. The foot is found distal to the ankle and contains 26 
bones. The ankle joint is formed by the articulations between the talus 
bone of the foot and the distal end of the tibia, the medial malleolus of 
the tibia, and the lateral malleolus of the fibula. The posterior foot 
contains the seven tarsal bones, which are the talus, calcaneus, 
navicular, cuboid, and the medial, intermediate, and lateral cuneiform 
bones. The anterior foot consists of the five metatarsal bones, which 
are numbered 1-5 starting on the medial side of the foot. The toes 
contain 14 phalanx bones, with the big toe (toe number 1) having a 
proximal and a distal phalanx, and the other toes having proximal, 
middle, and distal phalanges. 


Exercise: 


Problem: 


The talus bone of the foot receives the weight of the body from the 
tibia. The talus bone then distributes this weight toward the ground in 
two directions: one-half of the body weight is passed in a posterior 
direction and one-half of the weight is passed in an anterior direction. 
Describe the arrangement of the tarsal and metatarsal bones that are 
involved in both the posterior and anterior distribution of body weight. 


Solution: 


The talus bone articulates superiorly with the tibia and fibula at the 
ankle joint, with body weight passed from the tibia to the talus. Body 
weight from the talus is transmitted to the ground by both ends of the 
medial and lateral longitudinal foot arches. Weight is passed 
posteriorly through both arches to the calcaneus bone, which forms the 
heel of the foot and is in contact with the ground. On the medial side 
of the foot, body weight is passed anteriorly from the talus bone to the 
navicular bone, and then to the medial, intermediate, and lateral 
cuneiform bones. The cuneiform bones pass the weight anteriorly to 
the first, second, and third metatarsal bones, whose heads (distal ends) 
are in contact with the ground. On the lateral side, body weight is 
passed anteriorly from the talus through the calcaneus, cuboid, and 
fourth and fifth metatarsal bones. The talus bone thus transmits body 
weight posteriorly to the calcaneus and anteriorly through the 
navicular, cuneiform, and cuboid bones, and metatarsals one through 
five. 


Glossary 
adductor tubercle 
small, bony bump located on the superior aspect of the medial 


epicondyle of the femur 


ankle joint 


joint that separates the leg and foot portions of the lower limb; formed 
by the articulations between the talus bone of the foot inferiorly, and 
the distal end of the tibia, medial malleolus of the tibia, and lateral 
malleolus of the fibula superiorly 


anterior border of the tibia 
narrow, anterior margin of the tibia that extends inferiorly from the 
tibial tuberosity 


base of the metatarsal bone 
expanded, proximal end of each metatarsal bone 


calcaneus 
heel bone; posterior, inferior tarsal bone that forms the heel of the foot 


cuboid 
tarsal bone that articulates posteriorly with the calcaneus bone, 
medially with the lateral cuneiform bone, and anteriorly with the 
fourth and fifth metatarsal bones 


distal tibiofibular joint 
articulation between the distal fibula and the fibular notch of the tibia 


femur 
thigh bone; the single bone of the thigh 


fibula 
thin, non-weight-bearing bone found on the lateral side of the leg 


fibular notch 
wide groove on the lateral side of the distal tibia for articulation with 
the fibula at the distal tibiofibular joint 


foot 
portion of the lower limb located distal to the ankle joint 


fovea capitis 


minor indentation on the head of the femur that serves as the site of 
attachment for the ligament to the head of the femur 


gluteal tuberosity 
roughened area on the posterior side of the proximal femur, extending 
inferiorly from the base of the greater trochanter 


greater trochanter 
large, bony expansion of the femur that projects superiorly from the 
base of the femoral neck 


hallux 
big toe; digit 1 of the foot 


head of the femur 
rounded, proximal end of the femur that articulates with the 
acetabulum of the hip bone to form the hip joint 


head of the fibula 
small, knob-like, proximal end of the fibula; articulates with the 
inferior aspect of the lateral condyle of the tibia 


head of the metatarsal bone 
expanded, distal end of each metatarsal bone 


hip joint 
joint located at the proximal end of the lower limb; formed by the 
articulation between the acetabulum of the hip bone and the head of 
the femur 


intercondylar eminence 
irregular elevation on the superior end of the tibia, between the 
articulating surfaces of the medial and lateral condyles 


intercondylar fossa 
deep depression on the posterior side of the distal femur that separates 
the medial and lateral condyles 


intermediate cuneiform 
middle of the three cuneiform tarsal bones; articulates posteriorly with 
the navicular bone, medially with the medial cuneiform bone, laterally 
with the lateral cuneiform bone, and anteriorly with the second 
metatarsal bone 


interosseous border of the fibula 
small ridge running down the medial side of the fibular shaft; for 
attachment of the interosseous membrane between the fibula and tibia 


interosseous border of the tibia 
small ridge running down the lateral side of the tibial shaft; for 
attachment of the interosseous membrane between the tibia and fibula 


interosseous membrane of the leg 
sheet of dense connective tissue that unites the shafts of the tibia and 
fibula bones 


intertrochanteric crest 
short, prominent ridge running between the greater and lesser 
trochanters on the posterior side of the proximal femur 


intertrochanteric line 
small ridge running between the greater and lesser trochanters on the 
anterior side of the proximal femur 


knee joint 
joint that separates the thigh and leg portions of the lower limb; 
formed by the articulations between the medial and lateral condyles of 
the femur, and the medial and lateral condyles of the tibia 


lateral condyle of the femur 
smooth, articulating surface that forms the distal and posterior sides of 
the lateral expansion of the distal femur 


lateral condyle of the tibia 
lateral, expanded region of the proximal tibia that includes the smooth 
surface that articulates with the lateral condyle of the femur as part of 


the knee joint 


lateral cuneiform 
most lateral of the three cuneiform tarsal bones; articulates posteriorly 
with the navicular bone, medially with the intermediate cuneiform 
bone, laterally with the cuboid bone, and anteriorly with the third 
metatarsal bone 


lateral epicondyle of the femur 
roughened area of the femur located on the lateral side of the lateral 
condyle 


lateral malleolus 
expanded distal end of the fibula 


leg 
portion of the lower limb located between the knee and ankle joints 


lesser trochanter 
small, bony projection on the medial side of the proximal femur, at the 
base of the femoral neck 


ligament of the head of the femur 
ligament that spans the acetabulum of the hip bone and the fovea 
capitis of the femoral head 


linea aspera 
longitudinally running bony ridge located in the middle third of the 
posterior femur 


medial condyle of the femur 
smooth, articulating surface that forms the distal and posterior sides of 
the medial expansion of the distal femur 


medial condyle of the tibia 
medial, expanded region of the proximal tibia that includes the smooth 
surface that articulates with the medial condyle of the femur as part of 
the knee joint 


medial cuneiform 
most medial of the three cuneiform tarsal bones; articulates posteriorly 
with the navicular bone, laterally with the intermediate cuneiform 
bone, and anteriorly with the first and second metatarsal bones 


medial epicondyle of the femur 
roughened area of the distal femur located on the medial side of the 
medial condyle 


medial malleolus 
bony expansion located on the medial side of the distal tibia 


metatarsal bone 
one of the five elongated bones that forms the anterior half of the foot; 
numbered 1-5, starting on the medial side of the foot 


metatarsophalangeal joint 
articulation between a metatarsal bone of the foot and the proximal 
phalanx bone of a toe 


navicular 
tarsal bone that articulates posteriorly with the talus bone, laterally 
with the cuboid bone, and anteriorly with the medial, intermediate, and 
lateral cuneiform bones 


neck of the femur 
narrowed region located inferior to the head of the femur 


patella 
kneecap; the largest sesamoid bone of the body; articulates with the 
distal femur 


patellar surface 
smooth groove located on the anterior side of the distal femur, between 
the medial and lateral condyles; site of articulation for the patella 


phalanx bone of the foot 


(plural = phalanges) one of the 14 bones that form the toes; these 
include the proximal and distal phalanges of the big toe, and the 
proximal, middle, and distal phalanx bones of toes two through five 


proximal tibiofibular joint 
articulation between the head of the fibula and the inferior aspect of 
the lateral condyle of the tibia 


shaft of the femur 
cylindrically shaped region that forms the central portion of the femur 


shaft of the fibula 
elongated, slender portion located between the expanded ends of the 
fibula 


shaft of the tibia 
triangular-shaped, central portion of the tibia 


soleal line 
small, diagonally running ridge located on the posterior side of the 
proximal tibia 


sustentaculum tali 
bony ledge extending from the medial side of the calcaneus bone 


talus 
tarsal bone that articulates superiorly with the tibia and fibula at the 
ankle joint; also articulates inferiorly with the calcaneus bone and 
anteriorly with the navicular bone 


tarsal bone 
one of the seven bones that make up the posterior foot; includes the 
calcaneus, talus, navicular, cuboid, medial cuneiform, intermediate 
cuneiform, and lateral cuneiform bones 


thigh 
portion of the lower limb located between the hip and knee joints 


tibia 
shin bone; the large, weight-bearing bone located on the medial side of 
the leg 


tibial tuberosity 
elevated area on the anterior surface of the proximal tibia 


Classification of Joints 
By the end of this section, you will be able to: 


e Distinguish between the functional and structural classifications for 
joints 

¢ Describe the three functional types of joints and give an example of 
each 

e List the three types of diarthrodial joints 


A joint, also called an articulation, is any place where adjacent bones or 
bone and cartilage come together (articulate with each other) to form a 
connection. Joints are classified both structurally and functionally. 
Structural classifications of joints take into account whether the adjacent 
bones are strongly anchored to each other by fibrous connective tissue or 
cartilage, or whether the adjacent bones articulate with each other within a 
fluid-filled space called a joint cavity. Functional classifications describe 
the degree of movement available between the bones, ranging from 
immobile, to slightly mobile, to freely moveable joints. The amount of 
movement available at a particular joint of the body is related to the 
functional requirements for that joint. Thus immobile or slightly moveable 
joints serve to protect internal organs, give stability to the body, and allow 
for limited body movement. In contrast, freely moveable joints allow for 
much more extensive movements of the body and limbs. 


Structural Classification of Joints 


The structural classification of joints is based on whether the articulating 
surfaces of the adjacent bones are directly connected by fibrous connective 
tissue or cartilage, or whether the articulating surfaces contact each other 
within a fluid-filled joint cavity. These differences serve to divide the joints 
of the body into three structural classifications. A fibrous joint is where the 
adjacent bones are united by fibrous connective tissue. At a cartilaginous 
joint, the bones are joined by hyaline cartilage or fibrocartilage. At a 
synovial joint, the articulating surfaces of the bones are not directly 
connected, but instead come into contact with each other within a joint 
cavity that is filled with a lubricating fluid. Synovial joints allow for free 
movement between the bones and are the most common joints of the body. 


Functional Classification of Joints 


The functional classification of joints is determined by the amount of 
mobility found between the adjacent bones. Joints are thus functionally 
classified as a synarthrosis or immobile joint, an amphiarthrosis or slightly 
moveable joint, or as a diarthrosis, which is a freely moveable joint 
(arthroun = “to fasten by a joint”). Depending on their location, fibrous 
joints may be functionally classified as a synarthrosis (immobile joint) or an 
amphiarthrosis (slightly mobile joint). Cartilaginous joints are also 
functionally classified as either a synarthrosis or an amphiarthrosis joint. 
All synovial joints are functionally classified as a diarthrosis joint. 


Synarthrosis 


An immobile or nearly immobile joint is called a synarthrosis. The 
immobile nature of these joints provide for a strong union between the 
articulating bones. This is important at locations where the bones provide 
protection for internal organs. Examples include sutures, the fibrous joints 
between the bones of the skull that surround and protect the brain ([link]), 
and the manubriosternal joint, the cartilaginous joint that unites the 
manubrium and body of the sternum for protection of the heart. 

Suture Joints of Skull 


Coronal 
suture 


Lambdoid Squamous 
suture suture 


The suture joints of the skull are an 
example of a synarthrosis, an 
immobile or essentially immobile 
joint. 


Amphiarthrosis 


An amphiarthrosis is a joint that has limited mobility. An example of this 
type of joint is the cartilaginous joint that unites the bodies of adjacent 
vertebrae. Filling the gap between the vertebrae is a thick pad of 
fibrocartilage called an intervertebral disc ({link]). Each intervertebral disc 
strongly unites the vertebrae but still allows for a limited amount of 
movement between them. However, the small movements available 
between adjacent vertebrae can sum together along the length of the 
vertebral column to provide for large ranges of body movements. 


Another example of an amphiarthrosis is the pubic symphysis of the pelvis. 

This is a cartilaginous joint in which the pubic regions of the right and left 

hip bones are strongly anchored to each other by fibrocartilage. This joint 

normally has very little mobility. The strength of the pubic symphysis is 

important in conferring weight-bearing stability to the pelvis. 

Intervertebral Disc 
i 


Vertebral body 


Intervertebral disc 


Lateral view 


An intervertebral disc unites the 
bodies of adjacent vertebrae within 
the vertebral column. Each disc 
allows for limited movement 
between the vertebrae and thus 
functionally forms an 
amphiarthrosis type of joint. 
Intervertebral discs are made of 
fibrocartilage and thereby 
structurally form a symphysis type 
of cartilaginous joint. 


Diarthrosis 


A freely mobile joint is classified as a diarthrosis. These types of joints 
include all synovial joints of the body, which provide the majority of body 
movements. Most diarthrotic joints are found in the appendicular skeleton 
and thus give the limbs a wide range of motion. These joints are divided 
into three categories, based on the number of axes of motion provided by 
each. An axis in anatomy is described as the movements in reference to the 
three anatomical planes: transverse, frontal, and sagittal. Thus, diarthroses 
are Classified as uniaxial (for movement in one plane), biaxial (for 
movement in two planes), or multiaxial joints (for movement in all three 
anatomical planes). 


A uniaxial joint only allows for a motion in a single plane (around a single 
axis). The elbow joint, which only allows for bending or straightening, is an 
example of a uniaxial joint. A biaxial joint allows for motions within two 
planes. An example of a biaxial joint is a metacarpophalangeal joint 
(knuckle joint) of the hand. The joint allows for movement along one axis 
to produce bending or straightening of the finger, and movement along a 
second axis, which allows for spreading of the fingers away from each other 
and bringing them together. A joint that allows for the several directions of 
movement is called a multiaxial joint (polyaxial or triaxial joint). This type 


of diarthrotic joint allows for movement along three axes ([link]). The 
shoulder and hip joints are multiaxial joints. They allow the upper or lower 
limb to move in an anterior-posterior direction and a medial-lateral 
direction. In addition, the limb can also be rotated around its long axis. This 
third movement results in rotation of the limb so that its anterior surface is 
moved either toward or away from the midline of the body. 

Multiaxial Joint 


Acetabulum of 
hip bone 


Head of femur 


A multiaxial joint, such as the hip joint, allows for 
three types of movement: anterior-posterior, 
medial-lateral, and rotational. 


Chapter Review 


Structural classifications of the body joints are based on how the bones are 
held together and articulate with each other. At fibrous joints, the adjacent 
bones are directly united to each other by fibrous connective tissue. 
Similarly, at a cartilaginous joint, the adjacent bones are united by cartilage. 
In contrast, at a synovial joint, the articulating bone surfaces are not directly 
united to each other, but come together within a fluid-filled joint cavity. 


The functional classification of body joints is based on the degree of 
movement found at each joint. A synarthrosis is a joint that is essentially 


immobile. This type of joint provides for a strong connection between the 
adjacent bones, which serves to protect internal structures such as the brain 
or heart. Examples include the fibrous joints of the skull sutures and the 
cartilaginous manubriosternal joint. A joint that allows for limited 
movement is an amphiarthrosis. An example is the pubic symphysis of the 
pelvis, the cartilaginous joint that strongly unites the right and left hip bones 
of the pelvis. The cartilaginous joints in which vertebrae are united by 
intervertebral discs provide for small movements between the adjacent 
vertebrae and are also an amphiarthrosis type of joint. Thus, based on their 
movement ability, both fibrous and cartilaginous joints are functionally 
classified as a synarthrosis or amphiarthrosis. 


The most common type of joint is the diarthrosis, which is a freely 
moveable joint. All synovial joints are functionally classified as diarthroses. 
A uniaxial diarthrosis, such as the elbow, is a joint that only allows for 
movement within a single anatomical plane. Joints that allow for 
movements in two planes are biaxial joints, such as the 
metacarpophalangeal joints of the fingers. A multiaxial joint, such as the 
shoulder or hip joint, allows for three planes of motions. 


Review Questions 


Exercise: 


Problem: 


The joint between adjacent vertebrae that includes an invertebral disc 
is classified as which type of joint? 


a. diarthrosis 

b. multiaxial 

c. amphiarthrosis 
d. synarthrosis 


Solution: 


C 


Exercise: 


Problem: Which of these joints is classified as a synarthrosis? 


a. the pubic symphysis 

b. the manubriosternal joint 
c. an invertebral disc 

d. the shoulder joint 


Solution: 


B 


Exercise: 


Problem: Which of these joints is classified as a biaxial diarthrosis? 


a. the metacarpophalangeal joint 
b. the hip joint 

c. the elbow joint 

d. the pubic symphysis 


Solution: 


A 


Exercise: 


Problem:Synovial joints 


a. may be functionally classified as a synarthrosis 

b. are joints where the bones are connected to each other by hyaline 
cartilage 

c. may be functionally classified as a amphiarthrosis 

d. are joints where the bones articulate with each other within a 
fluid-filled joint cavity 


Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: 


Define how joints are classified based on function. Describe and give 
an example for each functional type of joint. 


Solution: 


Functional classification of joints is based on the degree of mobility 
exhibited by the joint. A synarthrosis is an immobile or nearly 
immobile joint. An example is the manubriosternal joint or the joints 
between the skull bones surrounding the brain. An amphiarthrosis is a 
slightly moveable joint, such as the pubic symphysis or an 
intervertebral cartilaginous joint. A diarthrosis is a freely moveable 
joint. These are subdivided into three categories. A uniaxial diarthrosis 
allows movement within a single anatomical plane or axis of motion. 
The elbow joint is an example. A biaxial diarthrosis, such as the 
metacarpophalangeal joint, allows for movement along two planes or 
axes. The hip and shoulder joints are examples of a multiaxial 
diarthrosis. These allow movements along three planes or axes. 


Exercise: 


Problem: 


Explain the reasons for why joints differ in their degree of mobility. 


Solution: 


The functional needs of joints vary and thus joints differ in their degree 
of mobility. A synarthrosis, which is an immobile joint, serves to 
strongly connect bones thus protecting internal organs such as the heart 
or brain. A slightly moveable amphiarthrosis provides for small 
movements, which in the vertebral column can add together to yield a 
much larger overall movement. The freedom of movement provided by 
a diarthrosis can allow for large movements, such as is seen with most 
joints of the limbs. 


Glossary 


amphiarthrosis 
slightly mobile joint 


articulation 
joint of the body 


biaxial joint 
type of diarthrosis; a joint that allows for movements within two 
planes (two axes) 


cartilaginous joint 
joint at which the bones are united by hyaline cartilage 
(synchondrosis) or fibrocartilage (symphysis) 


diarthrosis 
freely mobile joint 


fibrous joint 
joint where the articulating areas of the adjacent bones are connected 
by fibrous connective tissue 


joint 
site at which two or more bones or bone and cartilage come together 


(articulate) 


joint cavity 


space enclosed by the articular capsule of a synovial joint that is filled 
with synovial fluid and contains the articulating surfaces of the 
adjacent bones 


multiaxial joint 
type of diarthrosis; a joint that allows for movements within three 
planes (three axes) 


synarthrosis 
immobile or nearly immobile joint 


synovial joint 
joint at which the articulating surfaces of the bones are located within 
a joint cavity formed by an articular capsule 


uniaxial joint 


type of diarthrosis; joint that allows for motion within only one plane 
(one axis) 


Fibrous Joints 
By the end of this section, you will be able to: 


e Describe the structural features of fibrous joints 
e Distinguish between a suture, syndesmosis, and gomphosis 
e Give an example of each type of fibrous joint 


At a fibrous joint, the adjacent bones are directly connected to each other by 
fibrous connective tissue, and thus the bones do not have a joint cavity 
between them ({link]). The gap between the bones may be narrow or wide. 
There are three types of fibrous joints. A suture is the narrow fibrous joint 
found between most bones of the skull. At a syndesmosis joint, the bones 
are more widely separated but are held together by a narrow band of fibrous 
connective tissue called a ligament or a wide sheet of connective tissue 
called an interosseous membrane. This type of fibrous joint is found 
between the shaft regions of the long bones in the forearm and in the leg. 
Lastly, a gomphosis is the narrow fibrous joint between the roots of a tooth 
and the bony socket in the jaw into which the tooth fits. 

Fibrous Joints 


. 7 Suture line : 
Vv n e ~ 
ea . ~ 
\ ‘en, ) =D. 
| __ Suture = a | 
a Ulna ——— | | ; | 
= Ly Bhs Radius 
_ fx 
AS Syndesmosis 
= af v/\ Dense HT | Socket Root of 
i fibrous AniabRIGHER 7 Hie 
% connective /*ntebracnial | Genpheus 
~~ tissue Meena | \ 
j membrane ON 
ZN 


Periodontal 
ligament 


(a) (b) (c) 


Fibrous joints form strong connections between bones. (a) 
Sutures join most bones of the skull. (b) An interosseous 
membrane forms a syndesmosis between the radius and 

ulna bones of the forearm. (c) A gomphosis is a specialized 
fibrous joint that anchors a tooth to its socket in the jaw. 


Suture 


All the bones of the skull, except for the mandible, are joined to each other 
by a fibrous joint called a suture. The fibrous connective tissue found at a 
suture (“to bind or sew”) strongly unites the adjacent skull bones and thus 
helps to protect the brain and form the face. In adults, the skull bones are 
closely opposed and fibrous connective tissue fills the narrow gap between 
the bones. The suture is frequently convoluted, forming a tight union that 
prevents most movement between the bones. (See [link]a.) Thus, skull 
sutures are functionally classified as a synarthrosis, although some sutures 
may allow for slight movements between the cranial bones. 


In newborns and infants, the areas of connective tissue between the bones 
are much wider, especially in those areas on the top and sides of the skull 
that will become the sagittal, coronal, squamous, and lambdoid sutures. 
These broad areas of connective tissue are called fontanelles ((link]). 
During birth, the fontanelles provide flexibility to the skull, allowing the 
bones to push closer together or to overlap slightly, thus aiding movement 
of the infant’s head through the birth canal. After birth, these expanded 
regions of connective tissue allow for rapid growth of the skull and 
enlargement of the brain. The fontanelles greatly decrease in width during 
the first year after birth as the skull bones enlarge. When the connective 
tissue between the adjacent bones is reduced to a narrow layer, these fibrous 
joints are now called sutures. At some sutures, the connective tissue will 
ossify and be converted into bone, causing the adjacent bones to fuse to 
each other. This fusion between bones is called a synostosis (“joined by 
bone”). Examples of synostosis fusions between cranial bones are found 
both early and late in life. At the time of birth, the frontal and maxillary 
bones consist of right and left halves joined together by sutures, which 
disappear by the eighth year as the halves fuse together to form a single 
bone. Late in life, the sagittal, coronal, and lambdoid sutures of the skull 
will begin to ossify and fuse, causing the suture line to gradually disappear. 
The Newborn Skull 


Anterior fontanelle 


Parietal bone Frontal bone 


Ossification 
center 


Posterior 


fontanelle Sphenoidal 


fontanelle 


Mastoid 
fontanelle 


Occipital bone Temporal bone (squamous portion) 


Lateral view 


The fontanelles of a newborn’s skull are 
broad areas of fibrous connective tissue 
that form fibrous joints between the bones 
of the skull. 


Syndesmosis 


A syndesmosis (“fastened with a band”) is a type of fibrous joint in which 
two parallel bones are united to each other by fibrous connective tissue. The 
gap between the bones may be narrow, with the bones joined by ligaments, 
or the gap may be wide and filled in by a broad sheet of connective tissue 
called an interosseous membrane. 


In the forearm, the wide gap between the shaft portions of the radius and 
ulna bones are strongly united by an interosseous membrane (see [link]b). 
Similarly, in the leg, the shafts of the tibia and fibula are also united by an 
interosseous membrane. In addition, at the distal tibiofibular joint, the 
articulating surfaces of the bones lack cartilage and the narrow gap between 
the bones is anchored by fibrous connective tissue and ligaments on both 
the anterior and posterior aspects of the joint. Together, the interosseous 
membrane and these ligaments form the tibiofibular syndesmosis. 


The syndesmoses found in the forearm and leg serve to unite parallel bones 
and prevent their separation. However, a syndesmosis does not prevent all 
movement between the bones, and thus this type of fibrous joint is 
functionally classified as an amphiarthrosis. In the leg, the syndesmosis 
between the tibia and fibula strongly unites the bones, allows for little 
movement, and firmly locks the talus bone in place between the tibia and 
fibula at the ankle joint. This provides strength and stability to the leg and 
ankle, which are important during weight bearing. In the forearm, the 
interosseous membrane is flexible enough to allow for rotation of the radius 
bone during forearm movements. Thus in contrast to the stability provided 
by the tibiofibular syndesmosis, the flexibility of the antebrachial 
interosseous membrane allows for the much greater mobility of the forearm. 


The interosseous membranes of the leg and forearm also provide areas for 
muscle attachment. Damage to a syndesmotic joint, which usually results 
from a fracture of the bone with an accompanying tear of the interosseous 
membrane, will produce pain, loss of stability of the bones, and may 
damage the muscles attached to the interosseous membrane. If the fracture 
site is not properly immobilized with a cast or splint, contractile activity by 
these muscles can cause improper alignment of the broken bones during 
healing. 


Gomphosis 


A gomphosis (“fastened with bolts”) is the specialized fibrous joint that 
anchors the root of a tooth into its bony socket within the maxillary bone 
(upper jaw) or mandible bone (lower jaw) of the skull. A gomphosis is also 
known as a peg-and-socket joint. Spanning between the bony walls of the 
socket and the root of the tooth are numerous short bands of dense 
connective tissue, each of which is called a periodontal ligament (see 
[link]c). Due to the immobility of a gomphosis, this type of joint is 
functionally classified as a synarthrosis. 


Chapter Review 


Fibrous joints are where adjacent bones are strongly united by fibrous 
connective tissue. The gap filled by connective tissue may be narrow or 


wide. The three types of fibrous joints are sutures, gomphoses, and 
syndesmoses. A suture is the narrow fibrous joint that unites most bones of 
the skull. At a gomphosis, the root of a tooth is anchored across a narrow 
gap by periodontal ligaments to the walls of its socket in the bony jaw. A 
syndesmosis is the type of fibrous joint found between parallel bones. The 
gap between the bones may be wide and filled with a fibrous interosseous 
membrane, or it may narrow with ligaments spanning between the bones. 
Syndesmoses are found between the bones of the forearm (radius and ulna) 
and the leg (tibia and fibula). Fibrous joints strongly unite adjacent bones 
and thus serve to provide protection for internal organs, strength to body 
regions, or weight-bearing stability. 


Review Questions 


Exercise: 


Problem: Which type of fibrous joint connects the tibia and fibula? 


a. syndesmosis 
b. symphysis 
c. suture 

d. gomphosis 


Solution: 


A 


Exercise: 


Problem: An example of a wide fibrous joint is 


a. the interosseous membrane of the forearm 
b. a gomphosis 

c. a suture joint 

d. a synostosis 


Solution: 
A 
Exercise: 
Problem:A gomphosis 


a. is formed by an interosseous membrane 

b. connects the tibia and fibula bones of the leg 
c. contains a joint cavity 

d. anchors a tooth to the jaw 


Solution: 
D 
Exercise: 
Problem: A syndesmosis is 
a. a Narrow fibrous joint 
b. the type of joint that unites bones of the skull 


c. a fibrous joint that unites parallel bones 
d. the type of joint that anchors the teeth in the jaws 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: 


Distinguish between a narrow and wide fibrous joint and give an 
example of each. 


Solution: 


Narrow fibrous joints are found at a suture, gomphosis, or 
syndesmosis. A suture is the fibrous joint that joins the bones of the 
skull to each other (except the mandible). A gomphosis is the fibrous 
joint that anchors each tooth to its bony socket within the upper or 
lower jaw. The tooth is connected to the bony jaw by periodontal 
ligaments. A narrow syndesmosis is found at the distal tibiofibular 
joint where the bones are united by fibrous connective tissue and 
ligaments. A syndesmosis can also form a wide fibrous joint where the 
shafts of two parallel bones are connected by a broad interosseous 
membrane. The radius and ulna bones of the forearm and the tibia and 
fibula bones of the leg are united by interosseous membranes. 


Exercise: 


Problem: 


The periodontal ligaments are made of collagen fibers and are 
responsible for connecting the roots of the teeth to the jaws. Describe 
how scurvy, a disease that inhibits collagen production, can affect the 
teeth. 


Solution: 


The teeth are anchored into their sockets within the bony jaws by the 
periodontal ligaments. This is a gomphosis type of fibrous joint. In 
scurvy, collagen production is inhibited and the periodontal ligaments 
become weak. This will cause the teeth to become loose or even to fall 
out. 


Glossary 


fontanelles 
expanded areas of fibrous connective tissue that separate the braincase 
bones of the skull prior to birth and during the first year after birth 


gomphosis 
type of fibrous joint in which the root of a tooth is anchored into its 
bony jaw socket by strong periodontal ligaments 


interosseous membrane 
wide sheet of fibrous connective tissue that fills the gap between two 
parallel bones, forming a syndesmosis; found between the radius and 
ulna of the forearm and between the tibia and fibula of the leg 


ligament 
strong band of dense connective tissue spanning between bones 


periodontal ligament 
band of dense connective tissue that anchors the root of a tooth into the 
bony jaw socket 


suture 
fibrous joint that connects the bones of the skull (except the mandible); 
an immobile joint (synarthrosis) 


syndesmosis 
type of fibrous joint in which two separated, parallel bones are 
connected by an interosseous membrane 


synostosis 
site at which adjacent bones or bony components have fused together 


Cartilaginous Joints 
By the end of this section, you will be able to: 


e Describe the structural features of cartilaginous joints 
e Distinguish between a synchondrosis and symphysis 
e Give an example of each type of cartilaginous joint 


As the name indicates, at a cartilaginous joint, the adjacent bones are united 
by cartilage, a tough but flexible type of connective tissue. These types of 
joints lack a joint cavity and involve bones that are joined together by either 
hyaline cartilage or fibrocartilage ({link]). There are two types of 
cartilaginous joints. A synchondrosis is a cartilaginous joint where the 
bones are joined by hyaline cartilage. Also classified as a synchondrosis are 
places where bone is united to a cartilage structure, such as between the 
anterior end of a rib and the costal cartilage of the thoracic cage. The 
second type of cartilaginous joint is a symphysis, where the bones are 
joined by fibrocartilage. 

Cartiliginous Joints 


A 


(temporary 
hyaline 
cartilage joint) 


At cartilaginous joints, bones are united by hyaline 
cartilage to form a synchondrosis or by fibrocartilage to 
form a symphysis. (a) The hyaline cartilage of the 
epiphyseal plate (growth plate) forms a synchondrosis that 
unites the shaft (diaphysis) and end (epiphysis) of a long 
bone and allows the bone to grow in length. (b) The pubic 
portions of the right and left hip bones of the pelvis are 


joined together by fibrocartilage, forming the pubic 
symphysis. 


Synchondrosis 


A synchondrosis (“joined by cartilage’) is a cartilaginous joint where 
bones are joined together by hyaline cartilage, or where bone is united to 
hyaline cartilage. A synchondrosis may be temporary or permanent. A 
temporary synchondrosis is the epiphyseal plate (growth plate) of a growing 
long bone. The epiphyseal plate is the region of growing hyaline cartilage 
that unites the diaphysis (shaft) of the bone to the epiphysis (end of the 
bone). Bone lengthening involves growth of the epiphyseal plate cartilage 
and its replacement by bone, which adds to the diaphysis. For many years 
during childhood growth, the rates of cartilage growth and bone formation 
are equal and thus the epiphyseal plate does not change in overall thickness 
as the bone lengthens. During the late teens and early 20s, growth of the 
cartilage slows and eventually stops. The epiphyseal plate is then 
completely replaced by bone, and the diaphysis and epiphysis portions of 
the bone fuse together to form a single adult bone. This fusion of the 
diaphysis and epiphysis is a synostosis. Once this occurs, bone lengthening 
ceases. For this reason, the epiphyseal plate is considered to be a temporary 
synchondrosis. Because cartilage is softer than bone tissue, injury to a 
growing long bone can damage the epiphyseal plate cartilage, thus stopping 
bone growth and preventing additional bone lengthening. 


Growing layers of cartilage also form synchondroses that join together the 
ilium, ischium, and pubic portions of the hip bone during childhood and 
adolescence. When body growth stops, the cartilage disappears and is 
replaced by bone, forming synostoses and fusing the bony components 
together into the single hip bone of the adult. Similarly, synostoses unite the 
sacral vertebrae that fuse together to form the adult sacrum. 


Note: 


r 


Lares 


7 . 
openstax COLLEGE 
. 


Visit this website to view a radiograph (X-ray image) of a child’s hand and 
wrist. The growing bones of child have an epiphyseal plate that forms a 
synchondrosis between the shaft and end of a long bone. Being less dense 
than bone, the area of epiphyseal cartilage is seen on this radiograph as the 
dark epiphyseal gaps located near the ends of the long bones, including the 
radius, ulna, metacarpal, and phalanx bones. Which of the bones in this 
image do not show an epiphyseal plate (epiphyseal gap)? 


Examples of permanent synchondroses are found in the thoracic cage. One 
example is the first sternocostal joint, where the first rib is anchored to the 
manubrium by its costal cartilage. (The articulations of the remaining costal 
cartilages to the sternum are all synovial joints.) Additional synchondroses 
are formed where the anterior end of the other 11 ribs is joined to its costal 
cartilage. Unlike the temporary synchondroses of the epiphyseal plate, these 
permanent synchondroses retain their hyaline cartilage and thus do not 
ossify with age. Due to the lack of movement between the bone and 
cartilage, both temporary and permanent synchondroses are functionally 
classified as a synarthrosis. 


Symphysis 


A cartilaginous joint where the bones are joined by fibrocartilage is called a 
symphysis (“growing together”). Fibrocartilage is very strong because it 
contains numerous bundles of thick collagen fibers, thus giving it a much 
greater ability to resist pulling and bending forces when compared with 
hyaline cartilage. This gives symphyses the ability to strongly unite the 
adjacent bones, but can still allow for limited movement to occur. Thus, a 
symphysis is functionally classified as an amphiarthrosis. 


The gap separating the bones at a symphysis may be narrow or wide. 
Examples in which the gap between the bones is narrow include the pubic 
symphysis and the manubriosternal joint. At the pubic symphysis, the pubic 
portions of the right and left hip bones of the pelvis are joined together by 
fibrocartilage across a narrow gap. Similarly, at the manubriosternal joint, 
fibrocartilage unites the manubrium and body portions of the sternum. 


The intervertebral symphysis is a wide symphysis located between the 
bodies of adjacent vertebrae of the vertebral column. Here a thick pad of 
fibrocartilage called an intervertebral disc strongly unites the adjacent 
vertebrae by filling the gap between them. The width of the intervertebral 
symphysis is important because it allows for small movements between the 
adjacent vertebrae. In addition, the thick intervertebral disc provides 
cushioning between the vertebrae, which is important when carrying heavy 
objects or during high-impact activities such as running or jumping. 


Chapter Review 


There are two types of cartilaginous joints. A synchondrosis is formed 
when the adjacent bones are united by hyaline cartilage. A temporary 
synchondrosis is formed by the epiphyseal plate of a growing long bone, 
which is lost when the epiphyseal plate ossifies as the bone reaches 
maturity. The synchondrosis is thus replaced by a synostosis. Permanent 
synchondroses that do not ossify are found at the first sternocostal joint and 
between the anterior ends of the bony ribs and the junction with their costal 
cartilage. A symphysis is where the bones are joined by fibrocartilage and 
the gap between the bones may be narrow or wide. A narrow symphysis is 
found at the manubriosternal joint and at the pubic symphysis. A wide 
symphysis is the intervertebral symphysis in which the bodies of adjacent 
vertebrae are united by an intervertebral disc. 


Interactive Link Questions 


Exercise: 


Problem: 


Go to this website to view a radiograph (X-ray image) of a child’s 
hand and wrist. The growing bones of child have an epiphyseal plate 
that forms a synchondrosis between the shaft and end of a long bone. 
Being less dense than bone, the area of epiphyseal cartilage is seen on 
this radiograph as the dark epiphyseal gaps located near the ends of the 
long bones, including the radius, ulna, metacarpal, and phalanx bones. 
Which of the bones in this image do not show an epiphyseal plate 


(epiphyseal gap)? 


Solution: 
Although they are still growing, the carpal bones of the wrist area do 


not show an epiphyseal plate. Instead of elongating, these bones grow 
in diameter by adding new bone to their surfaces. 


Review Questions 


Exercise: 


Problem:A cartilaginous joint 


a. has a joint cavity 

b. is called a symphysis when the bones are united by fibrocartilage 
c. anchors the teeth to the jaws 

d. is formed by a wide sheet of fibrous connective tissue 


Solution: 


B 


Exercise: 


Problem: A synchondrosis is 


a. found at the pubic symphysis 

b. where bones are connected together with fibrocartilage 
c. a type of fibrous joint 

d. found at the first sternocostal joint of the thoracic cage 


Solution: 


D 


Exercise: 


Problem: Which of the following are joined by a symphysis? 


a. adjacent vertebrae 

b. the first rib and the sternum 

c. the end and shaft of a long bone 
d. the radius and ulna bones 


Solution: 


A 


Exercise: 


Problem: 


The epiphyseal plate of a growing long bone in a child is classified as 


a 


a. synchondrosis 
b. synostosis 

c. symphysis 

d. syndesmosis 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: 


Describe the two types of cartilaginous joints and give examples of 
each. 


Solution: 


Cartilaginous joints are where the adjacent bones are joined by 
cartilage. At a synchondrosis, the bones are united by hyaline cartilage. 
The epiphyseal plate of growing long bones and the first sternocostal 
joint that unites the first rib to the sternum are examples of 
synchondroses. At a symphysis, the bones are joined by fibrocartilage, 
which is strong and flexible. Symphysis joints include the 
intervertebral symphysis between adjacent vertebrae and the pubic 
symphysis that joins the pubic portions of the right and left hip bones. 


Exercise: 


Problem: 


Both functional and structural classifications can be used to describe 
an individual joint. Define the first sternocostal joint and the pubic 
symphysis using both functional and structural characteristics. 


Solution: 


The first sternocostal joint is a synchondrosis type of cartilaginous 
joint in which hyaline cartilage unites the first rib to the manubrium of 
the sternum. This forms an immobile (synarthrosis) type of joint. The 
pubic symphysis is a slightly mobile (amphiarthrosis) cartilaginous 
joint, where the pubic portions of the right and left hip bones are 
united by fibrocartilage, thus forming a symphysis. 


Glossary 


symphysis 
type of cartilaginous joint where the bones are joined by fibrocartilage 


synchondrosis 
type of cartilaginous joint where the bones are joined by hyaline 
cartilage 


Synovial Joints 
By the end of this section, you will be able to: 


¢ Describe the structural features of a synovial joint 

e Discuss the function of additional structures associated with synovial 
joints 

e List the six types of synovial joints and give an example of each 


Synovial joints are the most common type of joint in the body ((link]). A 
key structural characteristic for a synovial joint that is not seen at fibrous or 
cartilaginous joints is the presence of a joint cavity. This fluid-filled space is 
the site at which the articulating surfaces of the bones contact each other. 
Also unlike fibrous or cartilaginous joints, the articulating bone surfaces at 
a synovial joint are not directly connected to each other with fibrous 
connective tissue or cartilage. This gives the bones of a synovial joint the 
ability to move smoothly against each other, allowing for increased joint 
mobility. 

Synovial Joints 


Bone 


Synovial 
membrane 


Articular 
capsule 


Articular 
cartilage 


Joint cavity 
containing 


Bone synovial fluid 


Synovial joints allow for smooth 
movements between the adjacent bones. 
The joint is surrounded by an articular 
capsule that defines a joint cavity filled 
with synovial fluid. The articulating 
surfaces of the bones are covered by a 
thin layer of articular cartilage. Ligaments 
support the joint by holding the bones 
together and resisting excess or abnormal 
joint motions. 


Structural Features of Synovial Joints 


Synovial joints are characterized by the presence of a joint cavity. The walls 
of this space are formed by the articular capsule, a fibrous connective 
tissue structure that is attached to each bone just outside the area of the 
bone’s articulating surface. The bones of the joint articulate with each other 
within the joint cavity. 


Friction between the bones at a synovial joint is prevented by the presence 
of the articular cartilage, a thin layer of hyaline cartilage that covers the 
entire articulating surface of each bone. However, unlike at a cartilaginous 
joint, the articular cartilages of each bone are not continuous with each 
other. Instead, the articular cartilage acts like a Teflon® coating over the 
bone surface, allowing the articulating bones to move smoothly against 
each other without damaging the underlying bone tissue. Lining the inner 
surface of the articular capsule is a thin synovial membrane. The cells of 
this membrane secrete synovial fluid (synovia = “a thick fluid”), a thick, 
slimy fluid that provides lubrication to further reduce friction between the 
bones of the joint. This fluid also provides nourishment to the articular 
cartilage, which does not contain blood vessels. The ability of the bones to 
move smoothly against each other within the joint cavity, and the freedom 
of joint movement this provides, means that each synovial joint is 
functionally classified as a diarthrosis. 


Outside of their articulating surfaces, the bones are connected together by 
ligaments, which are strong bands of fibrous connective tissue. These 
strengthen and support the joint by anchoring the bones together and 
preventing their separation. Ligaments allow for normal movements at a 
joint, but limit the range of these motions, thus preventing excessive or 
abnormal joint movements. Ligaments are classified based on their 
relationship to the fibrous articular capsule. An extrinsic ligament is 
located outside of the articular capsule, an intrinsic ligament is fused to or 
incorporated into the wall of the articular capsule, and an intracapsular 
ligament is located inside of the articular capsule. 


At many synovial joints, additional support is provided by the muscles and 
their tendons that act across the joint. A tendon is the dense connective 
tissue structure that attaches a muscle to bone. As forces acting on a joint 
increase, the body will automatically increase the overall strength of 
contraction of the muscles crossing that joint, thus allowing the muscle and 
its tendon to serve as a “dynamic ligament” to resist forces and support the 
joint. This type of indirect support by muscles is very important at the 
shoulder joint, for example, where the ligaments are relatively weak. 


Additional Structures Associated with Synovial Joints 


A few synovial joints of the body have a fibrocartilage structure located 
between the articulating bones. This is called an articular disc, which is 
generally small and oval-shaped, or a meniscus, which is larger and C- 
shaped. These structures can serve several functions, depending on the 
specific joint. In some places, an articular disc may act to strongly unite the 
bones of the joint to each other. Examples of this include the articular discs 
found at the sternoclavicular joint or between the distal ends of the radius 
and ulna bones. At other synovial joints, the disc can provide shock 
absorption and cushioning between the bones, which is the function of each 
meniscus within the knee joint. Finally, an articular disc can serve to 
smooth the movements between the articulating bones, as seen at the 
temporomandibular joint. Some synovial joints also have a fat pad, which 
can serve as a cushion between the bones. 


Additional structures located outside of a synovial joint serve to prevent 
friction between the bones of the joint and the overlying muscle tendons or 
skin. A bursa (plural = bursae) is a thin connective tissue sac filled with 
lubricating liquid. They are located in regions where skin, ligaments, 
muscles, or muscle tendons can rub against each other, usually near a body 
joint ((link]). Bursae reduce friction by separating the adjacent structures, 
preventing them from rubbing directly against each other. Bursae are 
classified by their location. A subcutaneous bursa is located between the 
skin and an underlying bone. It allows skin to move smoothly over the 
bone. Examples include the prepatellar bursa located over the kneecap and 
the olecranon bursa at the tip of the elbow. A submuscular bursa is found 
between a muscle and an underlying bone, or between adjacent muscles. 
These prevent rubbing of the muscle during movements. A large 
submuscular bursa, the trochanteric bursa, is found at the lateral hip, 
between the greater trochanter of the femur and the overlying gluteus 
maximus muscle. A subtendinous bursa is found between a tendon and a 
bone. Examples include the subacromial bursa that protects the tendon of 
shoulder muscle as it passes under the acromion of the scapula, and the 
suprapatellar bursa that separates the tendon of the large anterior thigh 
muscle from the distal femur just above the knee. 

Bursae 


_\ | 


Posterior 
cruciate 
ligament 


Tendon of 
quadriceps 
femoris 


Suprapatellar 
bursa 


Patella 


Prepatellar 
bursa 


Anterior Synovial cavity 

reas Infrapatellar 

ligament 

2 fat pad 
Infrapatellar 

Tibia bursa 


Patellar 
ligament 


Bursae are fluid-filled sacs that serve to 
prevent friction between skin, muscle, or 


tendon and an underlying bone. Three 
major bursae and a fat pad are part of the 
complex joint that unites the femur and 
tibia of the leg. 


A tendon sheath is similar in structure to a bursa, but smaller. It is a 
connective tissue sac that surrounds a muscle tendon at places where the 
tendon crosses a joint. It contains a lubricating fluid that allows for smooth 
motions of the tendon during muscle contraction and joint movements. 


Note: 

Homeostatic Imbalances 

Bursitis 

Bursitis is the inflammation of a bursa near a joint. This will cause pain, 
swelling, or tenderness of the bursa and surrounding area, and may also 
result in joint stiffness. Bursitis is most commonly associated with the 
bursae found at or near the shoulder, hip, knee, or elbow joints. At the 
shoulder, subacromial bursitis may occur in the bursa that separates the 
acromion of the scapula from the tendon of a shoulder muscle as it passes 
deep to the acromion. In the hip region, trochanteric bursitis can occur in 
the bursa that overlies the greater trochanter of the femur, just below the 
lateral side of the hip. Ischial bursitis occurs in the bursa that separates the 
skin from the ischial tuberosity of the pelvis, the bony structure that is 
weight bearing when sitting. At the knee, inflammation and swelling of the 
bursa located between the skin and patella bone is prepatellar bursitis 
(“housemaid’s knee”), a condition more commonly seen today in roofers or 
floor and carpet installers who do not use knee pads. At the elbow, 
olecranon bursitis is inflammation of the bursa between the skin and 
olecranon process of the ulna. The olecranon forms the bony tip of the 
elbow, and bursitis here is also known as “student’s elbow.” 

Bursitis can be either acute (lasting only a few days) or chronic. It can arise 
from muscle overuse, trauma, excessive or prolonged pressure on the skin, 
rheumatoid arthritis, gout, or infection of the joint. Repeated acute 


episodes of bursitis can result in a chronic condition. Treatments for the 
disorder include antibiotics if the bursitis is caused by an infection, or anti- 
inflammatory agents, such as nonsteroidal anti-inflammatory drugs 
(NSAIDs) or corticosteroids if the bursitis is due to trauma or overuse. 
Chronic bursitis may require that fluid be drained, but additional surgery is 
usually not required. 


Types of Synovial Joints 


Synovial joints are subdivided based on the shapes of the articulating 
surfaces of the bones that form each joint. The six types of synovial joints 
are pivot, hinge, condyloid, saddle, plane, and ball-and socket-joints 
({link]). 

Types of Synovial Joints 


(f) Ball-and-socket joint 
(hip joint) 


(a) Pivot joint 
(between C1 and 
C2 vertebrae) 


=a 
T 


(b) Hinge joint 
(elbow) 


(e) Condyloid joint 
(between radius and 
carpal bones of wrist) 


(d) Plane joint 
(between tarsal bones) 


(c) Saddle joint 
(between trapezium 
carpal bone and 1st 
metacarpal bone) 


The six types of synovial joints allow the body to move in a variety 
of ways. (a) Pivot joints allow for rotation around an axis, such as 
between the first and second cervical vertebrae, which allows for 
side-to-side rotation of the head. (b) The hinge joint of the elbow 

works like a door hinge. (c) The articulation between the trapezium 


carpal bone and the first metacarpal bone at the base of the thumb is a 
saddle joint. (d) Plane joints, such as those between the tarsal bones 
of the foot, allow for limited gliding movements between bones. (e) 

The radiocarpal joint of the wrist is a condyloid joint. (f) The hip and 

shoulder joints are the only ball-and-socket joints of the body. 


Pivot Joint 


At a pivot joint, a rounded portion of a bone is enclosed within a ring 
formed partially by the articulation with another bone and partially by a 
ligament (see [link]a). The bone rotates within this ring. Since the rotation 
is around a single axis, pivot joints are functionally classified as a uniaxial 
diarthrosis type of joint. An example of a pivot joint is the atlantoaxial joint, 
found between the C1 (atlas) and C2 (axis) vertebrae. Here, the upward 
projecting dens of the axis articulates with the inner aspect of the atlas, 
where it is held in place by a ligament. Rotation at this joint allows you to 
turn your head from side to side. A second pivot joint is found at the 
proximal radioulnar joint. Here, the head of the radius is largely encircled 
by a ligament that holds it in place as it articulates with the radial notch of 
the ulna. Rotation of the radius allows for forearm movements. 


Hinge Joint 


In a hinge joint, the convex end of one bone articulates with the concave 
end of the adjoining bone (see [link]b). This type of joint allows only for 
bending and straightening motions along a single axis, and thus hinge joints 
are functionally classified as uniaxial joints. A good example is the elbow 
joint, with the articulation between the trochlea of the humerus and the 
trochlear notch of the ulna. Other hinge joints of the body include the knee, 
ankle, and interphalangeal joints between the phalanx bones of the fingers 
and toes. 


Condyloid Joint 


At a condyloid joint (ellipsoid joint), the shallow depression at the end of 
one bone articulates with a rounded structure from an adjacent bone or 
bones (see [link]e). The knuckle (metacarpophalangeal) joints of the hand 
between the distal end of a metacarpal bone and the proximal phalanx bone 
are condyloid joints. Another example is the radiocarpal joint of the wrist, 
between the shallow depression at the distal end of the radius bone and the 
rounded scaphoid, lunate, and triquetrum carpal bones. In this case, the 
articulation area has a more oval (elliptical) shape. Functionally, condyloid 
joints are biaxial joints that allow for two planes of movement. One 
movement involves the bending and straightening of the fingers or the 
anterior-posterior movements of the hand. The second movement is a side- 
to-side movement, which allows you to spread your fingers apart and bring 
them together, or to move your hand in a medial-going or lateral-going 
direction. 


Saddle Joint 


At a saddle joint, both of the articulating surfaces for the bones have a 
saddle shape, which is concave in one direction and convex in the other (see 
[link]c). This allows the two bones to fit together like a rider sitting on a 
saddle. Saddle joints are functionally classified as biaxial joints. The 
primary example is the first carpometacarpal joint, between the trapezium 
(a carpal bone) and the first metacarpal bone at the base of the thumb. This 
joint provides the thumb the ability to move away from the palm of the 
hand along two planes. Thus, the thumb can move within the same plane as 
the palm of the hand, or it can jut out anteriorly, perpendicular to the palm. 
This movement of the first carpometacarpal joint is what gives humans their 
distinctive “opposable” thumbs. The sternoclavicular joint is also classified 
as a Saddle joint. 


Plane Joint 


Ata plane joint (gliding joint), the articulating surfaces of the bones are 
flat or slightly curved and of approximately the same size, which allows the 
bones to slide against each other (see [link]d). The motion at this type of 
joint is usually small and tightly constrained by surrounding ligaments. 
Based only on their shape, plane joints can allow multiple movements, 
including rotation. Thus plane joints can be functionally classified as a 
multiaxial joint. However, not all of these movements are available to every 
plane joint due to limitations placed on it by ligaments or neighboring 
bones. Thus, depending upon the specific joint of the body, a plane joint 
may exhibit only a single type of movement or several movements. Plane 
joints are found between the carpal bones (intercarpal joints) of the wrist or 
tarsal bones (intertarsal joints) of the foot, between the clavicle and 
acromion of the scapula (acromioclavicular joint), and between the superior 
and inferior articular processes of adjacent vertebrae (zygapophysial joints). 


Ball-and-Socket Joint 


The joint with the greatest range of motion is the ball-and-socket joint. At 
these joints, the rounded head of one bone (the ball) fits into the concave 
articulation (the socket) of the adjacent bone (see [link ]f). The hip joint and 
the glenohumeral (shoulder) joint are the only ball-and-socket joints of the 
body. At the hip joint, the head of the femur articulates with the acetabulum 
of the hip bone, and at the shoulder joint, the head of the humerus 
articulates with the glenoid cavity of the scapula. 


Ball-and-socket joints are classified functionally as multiaxial joints. The 
femur and the humerus are able to move in both anterior-posterior and 
medial-lateral directions and they can also rotate around their long axis. The 
shallow socket formed by the glenoid cavity allows the shoulder joint an 
extensive range of motion. In contrast, the deep socket of the acetabulum 
and the strong supporting ligaments of the hip joint serve to constrain 
movements of the femur, reflecting the need for stability and weight- 
bearing ability at the hip. 


Note: 


Watch this video to see an animation of synovial joints in action. Synovial 
joints are places where bones articulate with each other inside of a joint 
cavity. The different types of synovial joints are the ball-and-socket joint 
(shoulder joint), hinge joint (knee), pivot joint (atlantoaxial joint, between 
C1 and C2 vertebrae of the neck), condyloid joint (radiocarpal joint of the 
wrist), saddle joint (first carpometacarpal joint, between the trapezium 
carpal bone and the first metacarpal bone, at the base of the thumb), and 
plane joint (facet joints of vertebral column, between superior and inferior 
articular processes). Which type of synovial joint allows for the widest 
range of motion? 


Note: 

Aging and the... 

Joints 

Arthritis is a common disorder of synovial joints that involves 
inflammation of the joint. This often results in significant joint pain, along 
with swelling, stiffness, and reduced joint mobility. There are more than 
100 different forms of arthritis. Arthritis may arise from aging, damage to 
the articular cartilage, autoimmune diseases, bacterial or viral infections, or 
unknown (probably genetic) causes. 

The most common type of arthritis is osteoarthritis, which is associated 
with aging and “wear and tear” of the articular cartilage ((link]). Risk 
factors that may lead to osteoarthritis later in life include injury to a joint; 
jobs that involve physical labor; sports with running, twisting, or throwing 
actions; and being overweight. These factors put stress on the articular 
cartilage that covers the surfaces of bones at synovial joints, causing the 
cartilage to gradually become thinner. As the articular cartilage layer wears 


down, more pressure is placed on the bones. The joint responds by 
increasing production of the lubricating synovial fluid, but this can lead to 
swelling of the joint cavity, causing pain and joint stiffness as the articular 
capsule is stretched. The bone tissue underlying the damaged articular 
cartilage also responds by thickening, producing irregularities and causing 
the articulating surface of the bone to become rough or bumpy. Joint 
movement then results in pain and inflammation. In its early stages, 
symptoms of osteoarthritis may be reduced by mild activity that “warms 
up” the joint, but the symptoms may worsen following exercise. In 
individuals with more advanced osteoarthritis, the affected joints can 
become more painful and therefore are difficult to use effectively, resulting 
in increased immobility. There is no cure for osteoarthritis, but several 
treatments can help alleviate the pain. Treatments may include lifestyle 
changes, such as weight loss and low-impact exercise, and over-the- 
counter or prescription medications that help to alleviate the pain and 
inflammation. For severe cases, joint replacement surgery (arthroplasty) 
may be required. 

Joint replacement is a very invasive procedure, so other treatments are 
always tried before surgery. However arthroplasty can provide relief from 
chronic pain and can enhance mobility within a few months following the 
surgery. This type of surgery involves replacing the articular surfaces of the 
bones with prosthesis (artificial components). For example, in hip 
arthroplasty, the worn or damaged parts of the hip joint, including the head 
and neck of the femur and the acetabulum of the pelvis, are removed and 
replaced with artificial joint components. The replacement head for the 
femur consists of a rounded ball attached to the end of a shaft that is 
inserted inside the diaphysis of the femur. The acetabulum of the pelvis is 
reshaped and a replacement socket is fitted into its place. The parts, which 
are always built in advance of the surgery, are sometimes custom made to 
produce the best possible fit for a patient. 

Gout is a form of arthritis that results from the deposition of uric acid 
crystals within a body joint. Usually only one or a few joints are affected, 
such as the big toe, knee, or ankle. The attack may only last a few days, but 
may return to the same or another joint. Gout occurs when the body makes 
too much uric acid or the kidneys do not properly excrete it. A diet with 
excessive fructose has been implicated in raising the chances of a 
susceptible individual developing gout. 


Other forms of arthritis are associated with various autoimmune diseases, 
bacterial infections of the joint, or unknown genetic causes. Autoimmune 
diseases, including rheumatoid arthritis, scleroderma, or systemic lupus 
erythematosus, produce arthritis because the immune system of the body 
attacks the body joints. In rheumatoid arthritis, the joint capsule and 
synovial membrane become inflamed. As the disease progresses, the 
articular cartilage is severely damaged or destroyed, resulting in joint 
deformation, loss of movement, and severe disability. The most commonly 
involved joints are the hands, feet, and cervical spine, with corresponding 
joints on both sides of the body usually affected, though not always to the 
same extent. Rheumatoid arthritis is also associated with lung fibrosis, 
vasculitis (inflammation of blood vessels), coronary heart disease, and 
premature mortality. With no known cure, treatments are aimed at 
alleviating symptoms. Exercise, anti-inflammatory and pain medications, 
various specific disease-modifying anti-rheumatic drugs, or surgery are 
used to treat rheumatoid arthritis. 


Osteoarthritis 
WL 


Decreased 


Normal hip joint joint space 


Exposed bone 


Worn 
cartilage 


Osteoarthritis of a synovial joint results from 
aging or prolonged joint wear and tear. These 
cause erosion and loss of the articular cartilage 
covering the surfaces of the bones, resulting in 
inflammation that causes joint stiffness and 
pain. 


Visit this website to learn about a patient who arrives at the hospital with 
joint pain and weakness in his legs. What caused this patient’s weakness? 


Rtg 


Watch this animation to observe hip replacement surgery (total hip 
arthroplasty), which can be used to alleviate the pain and loss of joint 
mobility associated with osteoarthritis of the hip joint. What is the most 
common cause of hip disability? 


Watch this video to learn about the symptoms and treatments for 
rheumatoid arthritis. Which system of the body malfunctions in rheumatoid 
arthritis and what does this cause? 


Chapter Review 


Synovial joints are the most common type of joints in the body. They are 
characterized by the presence of a joint cavity, inside of which the bones of 
the joint articulate with each other. The articulating surfaces of the bones at 
a synovial joint are not directly connected to each other by connective 
tissue or cartilage, which allows the bones to move freely against each 
other. The walls of the joint cavity are formed by the articular capsule. 
Friction between the bones is reduced by a thin layer of articular cartilage 
covering the surfaces of the bones, and by a lubricating synovial fluid, 
which is secreted by the synovial membrane. 


Synovial joints are strengthened by the presence of ligaments, which hold 
the bones together and resist excessive or abnormal movements of the joint. 
Ligaments are classified as extrinsic ligaments if they are located outside of 
the articular capsule, intrinsic ligaments if they are fused to the wall of the 
articular capsule, or intracapsular ligaments if they are located inside the 
articular capsule. Some synovial joints also have an articular disc 
(meniscus), which can provide padding between the bones, smooth their 
movements, or strongly join the bones together to strengthen the joint. 
Muscles and their tendons acting across a joint can also increase their 
contractile strength when needed, thus providing indirect support for the 
joint. 


Bursae contain a lubricating fluid that serves to reduce friction between 
structures. Subcutaneous bursae prevent friction between the skin and an 
underlying bone, submuscular bursae protect muscles from rubbing against 
a bone or another muscle, and a subtendinous bursa prevents friction 
between bone and a muscle tendon. Tendon sheaths contain a lubricating 
fluid and surround tendons to allow for smooth movement of the tendon as 
it crosses a joint. 


Based on the shape of the articulating bone surfaces and the types of 
movement allowed, synovial joints are classified into six types. At a pivot 
joint, one bone is held within a ring by a ligament and its articulation with a 
second bone. Pivot joints only allow for rotation around a single axis. These 
are found at the articulation between the C1 (atlas) and the dens of the C2 
(axis) vertebrae, which provides the side-to-side rotation of the head, or at 
the proximal radioulnar joint between the head of the radius and the radial 
notch of the ulna, which allows for rotation of the radius during forearm 
movements. Hinge joints, such as at the elbow, knee, ankle, or 
interphalangeal joints between phalanx bones of the fingers and toes, allow 
only for bending and straightening of the joint. Pivot and hinge joints are 
functionally classified as uniaxial joints. 


Condyloid joints are found where the shallow depression of one bone 
receives a rounded bony area formed by one or two bones. Condyloid joints 
are found at the base of the fingers (metacarpophalangeal joints) and at the 
wrist (radiocarpal joint). At a saddle joint, the articulating bones fit together 
like a rider and a saddle. An example is the first carpometacarpal joint 
located at the base of the thumb. Both condyloid and saddle joints are 
functionally classified as biaxial joints. 


Plane joints are formed between the small, flattened surfaces of adjacent 
bones. These joints allow the bones to slide or rotate against each other, but 
the range of motion is usually slight and tightly limited by ligaments or 
surrounding bones. This type of joint is found between the articular 
processes of adjacent vertebrae, at the acromioclavicular joint, or at the 
intercarpal joints of the hand and intertarsal joints of the foot. Ball-and- 
socket joints, in which the rounded head of a bone fits into a large 
depression or socket, are found at the shoulder and hip joints. Both plane 
and ball-and-sockets joints are classified functionally as multiaxial joints. 
However, ball-and-socket joints allow for large movements, while the 
motions between bones at a plane joint are small. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to see an animation of synovial joints in action. 
Synovial joints are places where bones articulate with each other 
inside of a joint cavity. The different types of synovial joints are the 
ball-and-socket joint (shoulder joint), hinge joint (knee), pivot joint 
(atlantoaxial joint, between C1 and C2 vertebrae of the neck), 
condyloid joint (radiocarpal joint of the wrist), saddle joint (first 
carpometacarpal joint, between the trapezium carpal bone and the first 
metacarpal bone, at the base of the thumb), and plane joint (facet joints 
of vertebral column, between superior and inferior articular processes). 
Which type of synovial joint allows for the widest ranges of motion? 


Solution: 


Ball-and-socket joint. 
Exercise: 
Problem: 
Visit this website to read about a patient who arrives at the hospital 


with joint pain and weakness in his legs. What caused this patient’s 
weakness? 


Solution: 


Gout is due to the accumulation of uric acid crystals in the body. 
Usually these accumulate within joints, causing joint pain. This patient 
also had crystals that accumulated in the space next to his spinal cord, 
thus compressing the spinal cord and causing muscle weakness. 


Exercise: 
Problem: 
Watch this animation to observe hip replacement surgery (total hip 
arthroplasty), which can be used to alleviate the pain and loss of joint 


mobility associated with osteoarthritis of the hip joint. What is the 
most common cause of hip disability? 


Solution: 


The most common cause of hip disability is osteoarthritis, a chronic 
disease in which the articular cartilage of the joint wears away, 
resulting in severe hip pain and stiffness. 


Exercise: 


Problem: 


Watch this video to learn about the symptoms and treatments for 
rheumatoid arthritis. Which system of the body malfunctions in 
rheumatoid arthritis and what does this cause? 


Solution: 


The immune system malfunctions and attacks healthy cells in the 
lining of your joints. This causes inflammation and pain in the joints 
and surrounding tissues. 


Review Questions 


Exercise: 


Problem: Which type of joint provides the greatest range of motion? 


a. ball-and-socket 
b. hinge 

c. condyloid 

d. plane 


Solution: 


A 


Exercise: 


Problem: Which type of joint allows for only uniaxial movement? 


a. saddle joint 

b. hinge joint 

c. condyloid joint 

d. ball-and-socket joint 


Solution: 


B 


Exercise: 


Problem: Which of the following is a type of synovial joint? 


a. a synostosis 

b. a suture 

c. a plane joint 

d. a synchondrosis 


Solution: 


C 


Exercise: 


Problem:A bursa 


a. surrounds a tendon at the point where the tendon crosses a joint 

b. secretes the lubricating fluid for a synovial joint 

c. prevents friction between skin and bone, or a muscle tendon and 
bone 

d. is the strong band of connective tissue that holds bones together at 
a synovial joint 


Solution: 


@ 


Exercise: 


Problem: At synovial joints, 


a. the articulating ends of the bones are directly connected by 
fibrous connective tissue 

b. the ends of the bones are enclosed within a space called a 
subcutaneous bursa 

c. intrinsic ligaments are located entirely inside of the articular 
capsule 

d. the joint cavity is filled with a thick, lubricating fluid 


Solution: 


a) 


Exercise: 


Problem: At a synovial joint, the synovial membrane 


a. forms the fibrous connective walls of the joint cavity 

b. is the layer of cartilage that covers the articulating surfaces of the 
bones 

c. forms the intracapsular ligaments 

d. secretes the lubricating synovial fluid 


Solution: 


D 


Exercise: 


Problem: Condyloid joints 


a. are a type of ball-and-socket joint 

b. include the radiocarpal joint 

c. are a uniaxial diarthrosis joint 

d. are found at the proximal radioulnar joint 


Solution: 
B 
Exercise: 
Problem:A meniscus is 


a. a fibrocartilage pad that provides padding between bones 

b. a fluid-filled space that prevents friction between a muscle tendon 
and underlying bone 

c. the articular cartilage that covers the ends of a bone at a synovial 
joint 

d. the lubricating fluid within a synovial joint 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: 


Describe the characteristic structures found at all synovial joints. 


Solution: 


All synovial joints have a joint cavity filled with synovial fluid that is 
the site at which the bones of the joint articulate with each other. The 


articulating surfaces of the bones are covered by articular cartilage, a 
thin layer of hyaline cartilage. The walls of the joint cavity are formed 
by the connective tissue of the articular capsule. The synovial 
membrane lines the interior surface of the joint cavity and secretes the 
synovial fluid. Synovial joints are directly supported by ligaments, 
which span between the bones of the joint. These may be located 
outside of the articular capsule (extrinsic ligaments), incorporated or 
fused to the wall of the articular capsule (intrinsic ligaments), or found 
inside of the articular capsule (intracapsular ligaments). Ligaments 
hold the bones together and also serve to resist or prevent excessive or 
abnormal movements of the joint. 


Exercise: 


Problem: 


Describe the structures that provide direct and indirect support for a 
synovial joint. 


Solution: 


Direct support for a synovial joint is provided by ligaments that 
strongly unite the bones of the joint and serve to resist excessive or 
abnormal movements. Some joints, such as the sternoclavicular joint, 
have an articular disc that is attached to both bones, where it provides 
direct support by holding the bones together. Indirect joint support is 
provided by the muscles and their tendons that act across a joint. 
Muscles will increase their contractile force to help support the joint 
by resisting forces acting on it. 


Glossary 

articular capsule 
connective tissue structure that encloses the joint cavity of a synovial 
joint 


articular cartilage 


thin layer of hyaline cartilage that covers the articulating surfaces of 
bones at a synovial joint 


articular disc 
meniscus; a fibrocartilage structure found between the bones of some 
synovial joints; provides padding or smooths movements between the 
bones; strongly unites the bones together 


ball-and-socket joint 
synovial joint formed between the spherical end of one bone (the ball) 
that fits into the depression of a second bone (the socket); found at the 
hip and shoulder joints; functionally classified as a multiaxial joint 


bursa 
connective tissue sac containing lubricating fluid that prevents friction 
between adjacent structures, such as skin and bone, tendons and bone, 
or between muscles 


condyloid joint 
synovial joint in which the shallow depression at the end of one bone 
receives a rounded end from a second bone or a rounded structure 
formed by two bones; found at the metacarpophalangeal joints of the 
fingers or the radiocarpal joint of the wrist; functionally classified as a 
biaxial joint 
extrinsic ligament 
ligament located outside of the articular capsule of a synovial joint 
hinge joint 
synovial joint at which the convex surface of one bone articulates with 


the concave surface of a second bone; includes the elbow, knee, ankle, 
and interphalangeal joints; functionally classified as a uniaxial joint 


intracapsular ligament 
ligament that is located within the articular capsule of a synovial joint 


intrinsic ligament 


ligament that is fused to or incorporated into the wall of the articular 
capsule of a synovial joint 


meniscus 
articular disc 


pivot joint 
synovial joint at which the rounded portion of a bone rotates within a 
ring formed by a ligament and an articulating bone; functionally 
classified as uniaxial joint 


plane joint 
synovial joint formed between the flattened articulating surfaces of 
adjacent bones; functionally classified as a multiaxial joint 


proximal radioulnar joint 
articulation between head of radius and radial notch of ulna; uniaxial 
pivot joint that allows for rotation of radius during 
pronation/supination of forearm 


saddle joint 
synovial joint in which the articulating ends of both bones are convex 
and concave in shape, such as at the first carpometacarpal joint at the 
base of the thumb; functionally classified as a biaxial joint 


subcutaneous bursa 
bursa that prevents friction between skin and an underlying bone 


submuscular bursa 
bursa that prevents friction between bone and a muscle or between 
adjacent muscles 


subtendinous bursa 
bursa that prevents friction between bone and a muscle tendon 


synovial fluid 
thick, lubricating fluid that fills the interior of a synovial joint 


synovial membrane 
thin layer that lines the inner surface of the joint cavity at a synovial 
joint; produces the synovial fluid 


tendon 
dense connective tissue structure that anchors a muscle to bone 


tendon sheath 
connective tissue that surrounds a tendon at places where the tendon 
crosses a joint; contains a lubricating fluid to prevent friction and 
allow smooth movements of the tendon 


Types of Body Movements 
By the end of this section, you will be able to: 


¢ Define the different types of body movements 
¢ Identify the joints that allow for these motions 


Synovial joints allow the body a tremendous range of movements. Each 
movement at a synovial joint results from the contraction or relaxation of 
the muscles that are attached to the bones on either side of the articulation. 
The type of movement that can be produced at a synovial joint is 
determined by its structural type. While the ball-and-socket joint gives the 
greatest range of movement at an individual joint, in other regions of the 
body, several joints may work together to produce a particular movement. 
Overall, each type of synovial joint is necessary to provide the body with its 
great flexibility and mobility. There are many types of movement that can 
occur at synovial joints ({link]). Movement types are generally paired, with 
one being the opposite of the other. Body movements are always described 
in relation to the anatomical position of the body: upright stance, with upper 
limbs to the side of body and palms facing forward. Refer to [link] as you 
go through this section. 


Note: 


meee OPENStAX COLLEGE 
— 


Watch this video to learn about anatomical motions. What motions involve 
increasing or decreasing the angle of the foot at the ankle? 


Movements of the Body, Part 1 


Extension 
Flexion Extension y = 


|} / 
f } 
€ 
——s# 


Extension 


(a) and (b) Angular movements: flexion and extension at the shoulder and knees (c) Angular movements: flexion and extension 
of the neck 


Extension 

S y 

WY 4) pf 
AS 


ie 
\s 
} 


iat | : yi 


\ / 
f A 
\ vad if 


Extension \ \ Flexion a4} 
— \\ ly f/ Lateral 
{ a, Un! | rotation 
: Adduction 
Medial 
rotation 
(d) Angular movements: flexion (e) Angular movements: abduction, adduction, (f) Rotation of the head, neck, and lower limb 
and extension of the vertebral and circumduction of the upper limb at the 
column shoulder 


Synovial joints give the body many ways in which to move. 
(a)—(b) Flexion and extension motions are in the sagittal 
(anterior—posterior) plane of motion. These movements take 
place at the shoulder, hip, elbow, knee, wrist, 
metacarpophalangeal, metatarsophalangeal, and 
interphalangeal joints. (c)-(d) Anterior bending of the head or 
vertebral column is flexion, while any posterior-going 
movement is extension. (e) Abduction and adduction are 


motions of the limbs, hand, fingers, or toes in the coronal 
(medial—lateral) plane of movement. Moving the limb or hand 
laterally away from the body, or spreading the fingers or toes, 
is abduction. Adduction brings the limb or hand toward or 
across the midline of the body, or brings the fingers or toes 
together. Circumduction is the movement of the limb, hand, or 
fingers in a circular pattern, using the sequential combination 
of flexion, adduction, extension, and abduction motions. 
Adduction/abduction and circumduction take place at the 
shoulder, hip, wrist, metacarpophalangeal, and 
metatarsophalangeal joints. (f) Turning of the head side to side 
or twisting of the body is rotation. Medial and lateral rotation 
of the upper limb at the shoulder or lower limb at the hip 
involves turning the anterior surface of the limb toward the 
midline of the body (medial or internal rotation) or away from 
the midline (lateral or external rotation). 


Movements of the Body, Part 2 


| \ 
| \ ee 
Pronation | \ Supination 
(Radius / | (radius and 
rotates / ulna are 
over ulna) parallel) 


Dorsiflexion 


\ 


= ani —\} Plantar flexion / , 
LU JM \ <a D>" Eversion 


(g) Pronation (P) and supination (S) (h) Dorsiflexion and plantar flexion (i) Inversion and eversion 


A 


Retraction Protraction 


Elevation of 
of mandible of mandible 


( | eae 


fs : Depression 
‘4 of mandible 


(j) Protraction and retraction (k) Elevation and depression (I) Opposition 


(g) Supination of the forearm turns the hand to the palm 
forward position in which the radius and ulna are parallel, 
while forearm pronation turns the hand to the palm 
backward position in which the radius crosses over the ulna 
to form an "X." (h) Dorsiflexion of the foot at the ankle 
joint moves the top of the foot toward the leg, while plantar 
flexion lifts the heel and points the toes. (i) Eversion of the 
foot moves the bottom (sole) of the foot away from the 
midline of the body, while foot inversion faces the sole 
toward the midline. (j) Protraction of the mandible pushes 
the chin forward, and retraction pulls the chin back. (k) 
Depression of the mandible opens the mouth, while 


elevation closes it. (1) Opposition of the thumb brings the 
tip of the thumb into contact with the tip of the fingers of 
the same hand and reposition brings the thumb back next to 
the index finger. 


Flexion and Extension 


Flexion and extension are movements that take place within the sagittal 
plane and involve anterior or posterior movements of the body or limbs. For 
the vertebral column, flexion (anterior flexion) is an anterior (forward) 
bending of the neck or body, while extension involves a posterior-directed 
motion, such as straightening from a flexed position or bending backward. 
Lateral flexion is the bending of the neck or body toward the right or left 
side. These movements of the vertebral column involve both the symphysis 
joint formed by each intervertebral disc, as well as the plane type of 
synovial joint formed between the inferior articular processes of one 
vertebra and the superior articular processes of the next lower vertebra. 


In the limbs, flexion decreases the angle between the bones (bending of the 
joint), while extension increases the angle and straightens the joint. For the 
upper limb, all anterior-going motions are flexion and all posterior-going 
motions are extension. These include anterior-posterior movements of the 
arm at the shoulder, the forearm at the elbow, the hand at the wrist, and the 
fingers at the metacarpophalangeal and interphalangeal joints. For the 
thumb, extension moves the thumb away from the palm of the hand, within 
the same plane as the palm, while flexion brings the thumb back against the 
index finger or into the palm. These motions take place at the first 
carpometacarpal joint. In the lower limb, bringing the thigh forward and 
upward is flexion at the hip joint, while any posterior-going motion of the 
thigh is extension. Note that extension of the thigh beyond the anatomical 
(standing) position is greatly limited by the ligaments that support the hip 
joint. Knee flexion is the bending of the knee to bring the foot toward the 
posterior thigh, and extension is the straightening of the knee. Flexion and 
extension movements are seen at the hinge, condyloid, saddle, and ball-and- 
socket joints of the limbs (see [link ]a-d). 


Hyperextension is the abnormal or excessive extension of a joint beyond 
its normal range of motion, thus resulting in injury. Similarly, hyperflexion 
is excessive flexion at a joint. Hyperextension injuries are common at hinge 
joints such as the knee or elbow. In cases of “whiplash” in which the head is 
suddenly moved backward and then forward, a patient may experience both 
hyperextension and hyperflexion of the cervical region. 


Abduction and Adduction 


Abduction and adduction motions occur within the coronal plane and 
involve medial-lateral motions of the limbs, fingers, toes, or thumb. 
Abduction moves the limb laterally away from the midline of the body, 
while adduction is the opposing movement that brings the limb toward the 
body or across the midline. For example, abduction is raising the arm at the 
shoulder joint, moving it laterally away from the body, while adduction 
brings the arm down to the side of the body. Similarly, abduction and 
adduction at the wrist moves the hand away from or toward the midline of 
the body. Spreading the fingers or toes apart is also abduction, while 
bringing the fingers or toes together is adduction. For the thumb, abduction 
is the anterior movement that brings the thumb to a 90° perpendicular 
position, pointing straight out from the palm. Adduction moves the thumb 
back to the anatomical position, next to the index finger. Abduction and 
adduction movements are seen at condyloid, saddle, and ball-and-socket 
joints (see [link]e). 


Circumduction 


Circumduction is the movement of a body region in a circular manner, in 
which one end of the body region being moved stays relatively stationary 
while the other end describes a circle. It involves the sequential 
combination of flexion, adduction, extension, and abduction at a joint. This 
type of motion is found at biaxial condyloid and saddle joints, and at 
multiaxial ball-and-sockets joints (see [link]e). 


Rotation 


Rotation can occur within the vertebral column, at a pivot joint, or at a 
ball-and-socket joint. Rotation of the neck or body is the twisting 
movement produced by the summation of the small rotational movements 
available between adjacent vertebrae. At a pivot joint, one bone rotates in 
relation to another bone. This is a uniaxial joint, and thus rotation is the 
only motion allowed at a pivot joint. For example, at the atlantoaxial joint, 
the first cervical (C1) vertebra (atlas) rotates around the dens, the upward 
projection from the second cervical (C2) vertebra (axis). This allows the 
head to rotate from side to side as when shaking the head “no.” The 
proximal radioulnar joint is a pivot joint formed by the head of the radius 
and its articulation with the ulna. This joint allows for the radius to rotate 
along its length during pronation and supination movements of the forearm. 


Rotation can also occur at the ball-and-socket joints of the shoulder and hip. 
Here, the humerus and femur rotate around their long axis, which moves the 
anterior surface of the arm or thigh either toward or away from the midline 
of the body. Movement that brings the anterior surface of the limb toward 
the midline of the body is called medial (internal) rotation. Conversely, 
rotation of the limb so that the anterior surface moves away from the 
midline is lateral (external) rotation (see [link |f). Be sure to distinguish 
medial and lateral rotation, which can only occur at the multiaxial shoulder 
and hip joints, from circumduction, which can occur at either biaxial or 
multiaxial joints. 


Supination and Pronation 


Supination and pronation are movements of the forearm. In the anatomical 
position, the upper limb is held next to the body with the palm facing 
forward. This is the supinated position of the forearm. In this position, the 
radius and ulna are parallel to each other. When the palm of the hand faces 
backward, the forearm is in the pronated position, and the radius and ulna 
form an X-shape. 


Supination and pronation are the movements of the forearm that go between 
these two positions. Pronation is the motion that moves the forearm from 
the supinated (anatomical) position to the pronated (palm backward) 
position. This motion is produced by rotation of the radius at the proximal 


radioulnar joint, accompanied by movement of the radius at the distal 
radioulnar joint. The proximal radioulnar joint is a pivot joint that allows 
for rotation of the head of the radius. Because of the slight curvature of the 
shaft of the radius, this rotation causes the distal end of the radius to cross 
over the distal ulna at the distal radioulnar joint. This crossing over brings 
the radius and ulna into an X-shape position. Supination is the opposite 
motion, in which rotation of the radius returns the bones to their parallel 
positions and moves the palm to the anterior facing (supinated) position. It 
helps to remember that supination is the motion you use when scooping up 
soup with a spoon (see [link]g). 


Dorsiflexion and Plantar Flexion 


Dorsiflexion and plantar flexion are movements at the ankle joint, which 
is a hinge joint. Lifting the front of the foot, so that the top of the foot 
moves toward the anterior leg is dorsiflexion, while lifting the heel of the 
foot from the ground or pointing the toes downward is plantar flexion. 
These are the only movements available at the ankle joint (see [link]h). 


Inversion and Eversion 


Inversion and eversion are complex movements that involve the multiple 
plane joints among the tarsal bones of the posterior foot (intertarsal joints) 
and thus are not motions that take place at the ankle joint. Inversion is the 
turning of the foot to angle the bottom of the foot toward the midline, while 
eversion turns the bottom of the foot away from the midline. The foot has a 
greater range of inversion than eversion motion. These are important 
motions that help to stabilize the foot when walking or running on an 
uneven surface and aid in the quick side-to-side changes in direction used 
during active sports such as basketball, racquetball, or soccer (see [link ]i). 


Protraction and Retraction 


Protraction and retraction are anterior-posterior movements of the scapula 
or mandible. Protraction of the scapula occurs when the shoulder is moved 
forward, as when pushing against something or throwing a ball. Retraction 


is the opposite motion, with the scapula being pulled posteriorly and 
medially, toward the vertebral column. For the mandible, protraction occurs 
when the lower jaw is pushed forward, to stick out the chin, while retraction 
pulls the lower jaw backward. (See [link]j.) 


Depression and Elevation 


Depression and elevation are downward and upward movements of the 
scapula or mandible. The upward movement of the scapula and shoulder is 
elevation, while a downward movement is depression. These movements 
are used to shrug your shoulders. Similarly, elevation of the mandible is the 
upward movement of the lower jaw used to close the mouth or bite on 
something, and depression is the downward movement that produces 
opening of the mouth (see [link ]k). 


Excursion 


Excursion is the side to side movement of the mandible. Lateral excursion 
moves the mandible away from the midline, toward either the right or left 
side. Medial excursion returns the mandible to its resting position at the 
midline. 


Superior Rotation and Inferior Rotation 


Superior and inferior rotation are movements of the scapula and are defined 
by the direction of movement of the glenoid cavity. These motions involve 
rotation of the scapula around a point inferior to the scapular spine and are 
produced by combinations of muscles acting on the scapula. During 
superior rotation, the glenoid cavity moves upward as the medial end of 
the scapular spine moves downward. This is a very important motion that 
contributes to upper limb abduction. Without superior rotation of the 
scapula, the greater tubercle of the humerus would hit the acromion of the 
scapula, thus preventing any abduction of the arm above shoulder height. 
Superior rotation of the scapula is thus required for full abduction of the 
upper limb. Superior rotation is also used without arm abduction when 
carrying a heavy load with your hand or on your shoulder. You can feel this 


rotation when you pick up a load, such as a heavy book bag and carry it on 
only one shoulder. To increase its weight-bearing support for the bag, the 
shoulder lifts as the scapula superiorly rotates. Inferior rotation occurs 
during limb adduction and involves the downward motion of the glenoid 
cavity with upward movement of the medial end of the scapular spine. 


Opposition and Reposition 


Opposition is the thumb movement that brings the tip of the thumb in 
contact with the tip of a finger. This movement is produced at the first 
carpometacarpal joint, which is a saddle joint formed between the 
trapezium carpal bone and the first metacarpal bone. Thumb opposition is 
produced by a combination of flexion and abduction of the thumb at this 
joint. Returning the thumb to its anatomical position next to the index finger 
is called reposition (see [link]l). 


Movements of the Joints 


Type of 
Joint Movement Example 
Atlantoaxial joint (C1- 
Pivot Uniaxial joint; allows C2 vertebrae 
rotational movement articulation); proximal 
radioulnar joint 
Uniaxial joint; allows Knee; elbow; ankle; 
Hinge flexion/extension interphalangeal joints 


movements of fingers and toes 


Movements of the Joints 


Type of 
Joint Movement 


Biaxial joint; allows 
flexion/extension, 

Condyloid abduction/adduction, and 
circumduction 
movements 


Biaxial joint; allows 
flexion/extension, 

Saddle abduction/adduction, and 
circumduction 
movements 


Multiaxial joint; allows 
inversion and eversion of 
foot, or flexion, 
extension, and lateral 
flexion of the vertebral 
column 


Plane 


Multiaxial joint; allows 
flexion/extension, 
Ball-and- abduction/adduction, 
socket circumduction, and 
medial/lateral rotation 
movements 


Chapter Review 


The variety of movements provided by the different types of synovial joints 


Example 


Metacarpophalangeal 
(knuckle) joints of 
fingers; radiocarpal 
joint of wrist; 
metatarsophalangeal 
joints for toes 


First carpometacarpal 
joint of the thumb; 
stemoclavicular joint 


Intertarsal joints of 
foot; superior-inferior 
articular process 
articulations between 
vertebrae 


Shoulder and hip joints 


allows for a large range of body motions and gives you tremendous 


mobility. These movements allow you to flex or extend your body or limbs, 
medially rotate and adduct your arms and flex your elbows to hold a heavy 
object against your chest, raise your arms above your head, rotate or shake 
your head, and bend to touch the toes (with or without bending your knees). 


Each of the different structural types of synovial joints also allow for 
specific motions. The atlantoaxial pivot joint provides side-to-side rotation 
of the head, while the proximal radioulnar articulation allows for rotation of 
the radius during pronation and supination of the forearm. Hinge joints, 
such as at the knee and elbow, allow only for flexion and extension. 
Similarly, the hinge joint of the ankle only allows for dorsiflexion and 
plantar flexion of the foot. 


Condyloid and saddle joints are biaxial. These allow for flexion and 
extension, and abduction and adduction. The sequential combination of 
flexion, adduction, extension, and abduction produces circumduction. 
Multiaxial plane joints provide for only small motions, but these can add 
together over several adjacent joints to produce body movement, such as 
inversion and eversion of the foot. Similarly, plane joints allow for flexion, 
extension, and lateral flexion movements of the vertebral column. The 
multiaxial ball and socket joints allow for flexion-extension, abduction- 
adduction, and circumduction. In addition, these also allow for medial 
(internal) and lateral (external) rotation. Ball-and-socket joints have the 
greatest range of motion of all synovial joints. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to learn about anatomical motions. What motions 
involve increasing or decreasing the angle of the foot at the ankle? 


Solution: 


Dorsiflexion of the foot at the ankle decreases the angle of the ankle 
joint, while plantar flexion increases the angle of the ankle joint. 


Chapter Review 


Exercise: 
Problem: 


The joints between the articular processes of adjacent vertebrae can 
contribute to which movement? 


a. lateral flexion 
b. circumduction 
c. dorsiflexion 

d. abduction 


Solution: 


A 
Exercise: 


Problem: 


Which motion moves the bottom of the foot away from the midline of 
the body? 


a. elevation 

b. dorsiflexion 

c. eversion 

d. plantar flexion 


Solution: 


GC 


Exercise: 


Problem: 


Movement of a body region in a circular movement at a condyloid 
joint is what type of motion? 


a. rotation 

b. elevation 

c. abduction 

d. circumduction 


Solution: 


D 


Exercise: 


Problem: Supination is the motion that moves the 


a. hand from the palm backward position to the palm forward 
position 

b. foot so that the bottom of the foot faces the midline of the body 

c. hand from the palm forward position to the palm backward 
position 

d. scapula in an upward direction 


Solution: 


A 
Exercise: 


Problem: 


Movement at the shoulder joint that moves the upper limb laterally 
away from the body is called 


a. elevation 


b. eversion 
c. abduction 
d. lateral rotation 


Solution: 


‘Ss 


Critical Thinking Questions 


Exercise: 
Problem: 


Briefly define the types of joint movements available at a ball-and- 
socket joint. 


Solution: 


Ball-and-socket joints are multiaxial joints that allow for flexion and 
extension, abduction and adduction, circumduction, and medial and 
lateral rotation. 


Exercise: 


Problem: 


Discuss the joints involved and movements required for you to cross 
your arms together in front of your chest. 


Solution: 


To cross your arms, you need to use both your shoulder and elbow 
joints. At the shoulder, the arm would need to flex and medially rotate. 
At the elbow, the forearm would need to be flexed. 


Glossary 


abduction 
movement in the coronal plane that moves a limb laterally away from 
the body; spreading of the fingers 


adduction 
movement in the coronal plane that moves a limb medially toward or 
across the midline of the body; bringing fingers together 


circumduction 
circular motion of the arm, thigh, hand, thumb, or finger that is 
produced by the sequential combination of flexion, abduction, 
extension, and adduction 


depression 
downward (inferior) motion of the scapula or mandible 


dorsiflexion 
movement at the ankle that brings the top of the foot toward the 
anterior leg 


elevation 
upward (superior) motion of the scapula or mandible 


eversion 
foot movement involving the intertarsal joints of the foot in which the 
bottom of the foot is turned laterally, away from the midline 


extension 
movement in the sagittal plane that increases the angle of a joint 
(straightens the joint); motion involving posterior bending of the 
vertebral column or returning to the upright position from a flexed 
position 


flexion 
movement in the sagittal plane that decreases the angle of a joint 
(bends the joint); motion involving anterior bending of the vertebral 
column 


hyperextension 
excessive extension of joint, beyond the normal range of movement 


hyperflexion 
excessive flexion of joint, beyond the normal range of movement 


inferior rotation 
movement of the scapula during upper limb adduction in which the 
glenoid cavity of the scapula moves in a downward direction as the 
medial end of the scapular spine moves in an upward direction 


inversion 
foot movement involving the intertarsal joints of the foot in which the 
bottom of the foot is turned toward the midline 


lateral excursion 
side-to-side movement of the mandible away from the midline, toward 
either the right or left side 


lateral flexion 
bending of the neck or body toward the right or left side 


lateral (external) rotation 
movement of the arm at the shoulder joint or the thigh at the hip joint 
that moves the anterior surface of the limb away from the midline of 
the body 


medial excursion 
side-to-side movement that returns the mandible to the midline 


medial (internal) rotation 
movement of the arm at the shoulder joint or the thigh at the hip joint 
that brings the anterior surface of the limb toward the midline of the 
body 


opposition 
thumb movement that brings the tip of the thumb in contact with the 
tip of a finger 


plantar flexion 
foot movement at the ankle in which the heel is lifted off of the ground 


pronated position 
forearm position in which the palm faces backward 


pronation 
forearm motion that moves the palm of the hand from the palm 
forward to the palm backward position 


protraction 
anterior motion of the scapula or mandible 


reposition 
movement of the thumb from opposition back to the anatomical 
position (next to index finger) 


retraction 
posterior motion of the scapula or mandible 


rotation 
movement of a bone around a central axis (atlantoaxial joint) or around 
its long axis (proximal radioulnar joint; shoulder or hip joint); twisting 
of the vertebral column resulting from the summation of small motions 
between adjacent vertebrae 


superior rotation 
movement of the scapula during upper limb abduction in which the 
glenoid cavity of the scapula moves in an upward direction as the 
medial end of the scapular spine moves in a downward direction 


supinated position 
forearm position in which the palm faces anteriorly (anatomical 
position) 


supination 
forearm motion that moves the palm of the hand from the palm 
backward to the palm forward position 


Anatomy of Selected Synovial Joints 
By the end of this section, you will be able to: 


e Describe the bones that articulate together to form selected synovial 
joints 

e Discuss the movements available at each joint 

e Describe the structures that support and prevent excess movements at 
each joint 


Each synovial joint of the body is specialized to perform certain 
movements. The movements that are allowed are determined by the 
structural classification for each joint. For example, a multiaxial ball-and- 
socket joint has much more mobility than a uniaxial hinge joint. However, 
the ligaments and muscles that support a joint may place restrictions on the 
total range of motion available. Thus, the ball-and-socket joint of the 
shoulder has little in the way of ligament support, which gives the shoulder 
a very large range of motion. In contrast, movements at the hip joint are 
restricted by strong ligaments, which reduce its range of motion but confer 
stability during standing and weight bearing. 


This section will examine the anatomy of selected synovial joints of the 
body. Anatomical names for most joints are derived from the names of the 
bones that articulate at that joint, although some joints, such as the elbow, 
hip, and knee joints are exceptions to this general naming scheme. 


Articulations of the Vertebral Column 


In addition to being held together by the intervertebral discs, adjacent 
vertebrae also articulate with each other at synovial joints formed between 
the superior and inferior articular processes called zygapophysial joints 
(facet joints) (see [link]). These are plane joints that provide for only 
limited motions between the vertebrae. The orientation of the articular 
processes at these joints varies in different regions of the vertebral column 
and serves to determine the types of motions available in each vertebral 
region. The cervical and lumbar regions have the greatest ranges of 
motions. 


In the neck, the articular processes of cervical vertebrae are flattened and 
generally face upward or downward. This orientation provides the cervical 
vertebral column with extensive ranges of motion for flexion, extension, 
lateral flexion, and rotation. In the thoracic region, the downward projecting 
and overlapping spinous processes, along with the attached thoracic cage, 
greatly limit flexion, extension, and lateral flexion. However, the flattened 
and vertically positioned thoracic articular processes allow for the greatest 
range of rotation within the vertebral column. The lumbar region allows for 
considerable extension, flexion, and lateral flexion, but the orientation of 
the articular processes largely prohibits rotation. 


The articulations formed between the skull, the atlas (C1 vertebra), and the 
axis (C2 vertebra) differ from the articulations in other vertebral areas and 
play important roles in movement of the head. The atlanto-occipital joint 
is formed by the articulations between the superior articular processes of the 
atlas and the occipital condyles on the base of the skull. This articulation 
has a pronounced U-shaped curvature, oriented along the anterior-posterior 
axis. This allows the skull to rock forward and backward, producing flexion 
and extension of the head. This moves the head up and down, as when 
shaking your head “yes.” 


The atlantoaxial joint, between the atlas and axis, consists of three 
articulations. The paired superior articular processes of the axis articulate 
with the inferior articular processes of the atlas. These articulating surfaces 
are relatively flat and oriented horizontally. The third articulation is the 
pivot joint formed between the dens, which projects upward from the body 
of the axis, and the inner aspect of the anterior arch of the atlas ({link]). A 
strong ligament passes posterior to the dens to hold it in position against the 
anterior arch. These articulations allow the atlas to rotate on top of the axis, 
moving the head toward the right or left, as when shaking your head “no.” 
Atlantoaxial Joint 


Dens of 
C2 (axis) 


Anterior arch 


Superior articular of C1 (atlas) 


facet 


Ligament 


Superior view of atlas 


The atlantoaxial joint is a pivot type 
of joint between the dens portion of 
the axis (C2 vertebra) and the anterior 
arch of the atlas (C1 vertebra), with 
the dens held in place by a ligament. 


Temporomandibular Joint 


The temporomandibular joint (TMJ) is the joint that allows for opening 
(mandibular depression) and closing (mandibular elevation) of the mouth, 
as well as side-to-side and protraction/retraction motions of the lower jaw. 
This joint involves the articulation between the mandibular fossa and 
articular tubercle of the temporal bone, with the condyle (head) of the 
mandible. Located between these bony structures, filling the gap between 
the skull and mandible, is a flexible articular disc ([link]). This disc serves 
to smooth the movements between the temporal bone and mandibular 
condyle. 


Movement at the TMJ during opening and closing of the mouth involves 
both gliding and hinge motions of the mandible. With the mouth closed, the 
mandibular condyle and articular disc are located within the mandibular 
fossa of the temporal bone. During opening of the mouth, the mandible 
hinges downward and at the same time is pulled anteriorly, causing both the 
condyle and the articular disc to glide forward from the mandibular fossa 


onto the downward projecting articular tubercle. The net result is a forward 
and downward motion of the condyle and mandibular depression. The 
temporomandibular joint is supported by an extrinsic ligament that anchors 
the mandible to the skull. This ligament spans the distance between the base 
of the skull and the lingula on the medial side of the mandibular ramus. 


Dislocation of the TMJ may occur when opening the mouth too wide (such 
as when taking a large bite) or following a blow to the jaw, resulting in the 
mandibular condyle moving beyond (anterior to) the articular tubercle. In 
this case, the individual would not be able to close his or her mouth. 
Temporomandibular joint disorder is a painful condition that may arise due 
to arthritis, wearing of the articular cartilage covering the bony surfaces of 
the joint, muscle fatigue from overuse or grinding of the teeth, damage to 
the articular disc within the joint, or jaw injury. Temporomandibular joint 
disorders can also cause headache, difficulty chewing, or even the inability 
to move the jaw (lock jaw). Pharmacologic agents for pain or other 
therapies, including bite guards, are used as treatments. 
Temporomandibular Joint 


Articular disc 


Mandibular 
fossa 
uperior SOT aay rticular 
joint cay tubercle 
a 
Interior ——_$ —————— 
joint cavity ; Articular 
capsule 
Mandibular 
condyle 
Ramus of 
mandible 


The temporomandibular joint is the 
articulation between the temporal bone of 
the skull and the condyle of the mandible, 


with an articular disc located between 
these bones. During depression of the 
mandible (opening of the mouth), the 
mandibular condyle moves both forward 
and hinges downward as it travels from 
the mandibular fossa onto the articular 
tubercle. 


a 


gC 


Watch this video to learn about TMJ. Opening of the mouth requires the 
combination of two motions at the temporomandibular joint, an anterior 
gliding motion of the articular disc and mandible and the downward 
hinging of the mandible. What is the initial movement of the mandible 
during opening and how much mouth opening does this produce? 


Shoulder Joint 


The shoulder joint is called the glenohumeral joint. This is a ball-and- 
socket joint formed by the articulation between the head of the humerus and 
the glenoid cavity of the scapula ([link]). This joint has the largest range of 
motion of any joint in the body. However, this freedom of movement is due 


to the lack of structural support and thus the enhanced mobility is offset by 
a loss of stability. 


Glenohumeral Joint 


Clavicle Acromioclavicular ligament 


Tendon of 
supraspinatus 
muscle —— 


Glenoid labrum 


Acromion of scapula 


= Coracoacromial 
FBT ligament 

; eel Subacromial bursa 
Glenoid cavity SSRs 

Scapula ———__ “s= 5 


~~ 


Articular capsule 


Tendon sheath 


Tendon of biceps 
brachii muscles 


Articular cartilage Q\ . 
7 (long heed) 


Articular capsule: 
Synovial membrane 
Fibrous membrane 


Head of humerus 


ml Humerus 


The glenohumeral (shoulder) joint is a ball-and- 
socket joint that provides the widest range of 
motions. It has a loose articular capsule and is 
supported by ligaments and the rotator cuff 
muscles. 


The large range of motions at the shoulder joint is provided by the 
articulation of the large, rounded humeral head with the small and shallow 
glenoid cavity, which is only about one third of the size of the humeral 
head. The socket formed by the glenoid cavity is deepened slightly by a 
small lip of fibrocartilage called the glenoid labrum, which extends around 
the outer margin of the cavity. The articular capsule that surrounds the 
glenohumeral joint is relatively thin and loose to allow for large motions of 
the upper limb. Some structural support for the joint is provided by 
thickenings of the articular capsule wall that form weak intrinsic ligaments. 
These include the coracohumeral ligament, running from the coracoid 
process of the scapula to the anterior humerus, and three ligaments, each 
called a glenohumeral ligament, located on the anterior side of the 
articular capsule. These ligaments help to strengthen the superior and 
anterior capsule walls. 


However, the primary support for the shoulder joint is provided by muscles 
crossing the joint, particularly the four rotator cuff muscles. These muscles 
(supraspinatus, infraspinatus, teres minor, and subscapularis) arise from the 
scapula and attach to the greater or lesser tubercles of the humerus. As these 
muscles cross the shoulder joint, their tendons encircle the head of the 
humerus and become fused to the anterior, superior, and posterior walls of 
the articular capsule. The thickening of the capsule formed by the fusion of 
these four muscle tendons is called the rotator cuff. Two bursae, the 
subacromial bursa and the subscapular bursa, help to prevent friction 
between the rotator cuff muscle tendons and the scapula as these tendons 
cross the glenohumeral joint. In addition to their individual actions of 
moving the upper limb, the rotator cuff muscles also serve to hold the head 
of the humerus in position within the glenoid cavity. By constantly 
adjusting their strength of contraction to resist forces acting on the shoulder, 
these muscles serve as “dynamic ligaments” and thus provide the primary 
structural support for the glenohumeral joint. 


Injuries to the shoulder joint are common. Repetitive use of the upper limb, 
particularly in abduction such as during throwing, swimming, or racquet 
sports, may lead to acute or chronic inflammation of the bursa or muscle 
tendons, a tear of the glenoid labrum, or degeneration or tears of the rotator 
cuff. Because the humeral head is strongly supported by muscles and 
ligaments around its anterior, superior, and posterior aspects, most 
dislocations of the humerus occur in an inferior direction. This can occur 
when force is applied to the humerus when the upper limb is fully abducted, 
as when diving to catch a baseball and landing on your hand or elbow. 
Inflammatory responses to any shoulder injury can lead to the formation of 
scar tissue between the articular capsule and surrounding structures, thus 
reducing shoulder mobility, a condition called adhesive capsulitis (“frozen 
shoulder”). 


Note: 


Dea a 
Pie 


— 
= Shenatax 6 COLLEGE 


Watch this video for a tutorial on the anatomy of the shoulder joint. What 
movements are available at the shoulder joint? 


Note: 


openstax COLLEGE 


Obs fal) 


Watch this video to learn more about the anatomy of the shoulder joint, 
including bones, joints, muscles, nerves, and blood vessels. What is the 
shape of the glenoid labrum in cross-section, and what is the importance of 
this shape? 


Elbow Joint 


The elbow joint is a uniaxial hinge joint formed by the humeroulnar joint, 
the articulation between the trochlea of the humerus and the trochlear notch 
of the ulna. Also associated with the elbow are the humeroradial joint and 
the proximal radioulnar joint. All three of these joints are enclosed within a 
single articular capsule ([link]). 


The articular capsule of the elbow is thin on its anterior and posterior 
aspects, but is thickened along its outside margins by strong intrinsic 
ligaments. These ligaments prevent side-to-side movements and 

hyperextension. On the medial side is the triangular ulnar collateral 


ligament. This arises from the medial epicondyle of the humerus and 
attaches to the medial side of the proximal ulna. The strongest part of this 
ligament is the anterior portion, which resists hyperextension of the elbow. 
The ulnar collateral ligament may be injured by frequent, forceful 
extensions of the forearm, as is seen in baseball pitchers. Reconstructive 
surgical repair of this ligament is referred to as Tommy John surgery, named 
for the former major league pitcher who was the first person to have this 
treatment. 


The lateral side of the elbow is supported by the radial collateral ligament. 
This arises from the lateral epicondyle of the humerus and then blends into 
the lateral side of the annular ligament. The annular ligament encircles the 
head of the radius. This ligament supports the head of the radius as it 
articulates with the radial notch of the ulna at the proximal radioulnar joint. 
This is a pivot joint that allows for rotation of the radius during supination 
and pronation of the forearm. 

Elbow Joint 


Humerus Articular 


capsule 


Fat pad 


Synovial 
membrane 
Tendon ; c Synovial 
of triceps 7 - - cavity 
muscle \F) Articular 
i. cartilage 
Bursa | \\ of trochlea 
i. \ } ) fj = Tendon of 
SY . + 
Trochlea wi branchialis 
By — . muscle 
Articular } 


cartilage of the 
trochlear notch 


Olecranon bursa 


| 

| eve ' . NX Una 
al Coronoid 

= process 


(a) Medial sagittal section through right elbow (lateral view) 
Articular capsule 


oF Anular ligament 


\ : a —— 
ee Radius 
» 
Ulnar \ = = aa 
collateral —Lil ; D, ~— 
ligament AW ) “aa 
k TU) \\\ __ Una 
, Cael 
a 


Coronoid process 


(c) Medial view of right elbow joint 


Humerus j | 
ie 
\ Lateral epicondyle 
\ a ae Ulna 
Articular | =a jn ular 
capsule Min : ade 
p / tT Radial 
collateral 
\ 


Radius 


\ ligament 
—_ aw 
Olecranon ye = 
process 


(b) Lateral view of right elbow joint 


(a) The elbow is a hinge joint that allows only for flexion and 
extension of the forearm. (b) It is supported by the ulnar and radial 
collateral ligaments. (c) The annular ligament supports the head of the 
radius at the proximal radioulnar joint, the pivot joint that allows for 
rotation of the radius. 


Note: 


openstax COLLEGE” 


= “A 
1 
eye 


Watch this animation to learn more about the anatomy of the elbow joint. 
Which structures provide the main stability for the elbow? 


Note: 


ORR pao 
a r 


—s 
——. 
meee OPENStAX COLLEGE 


“ 
1 


eal 


Watch this video to learn more about the anatomy of the elbow joint, 
including bones, joints, muscles, nerves, and blood vessels. What are the 
functions of the articular cartilage? 


Hip Joint 


The hip joint is a multiaxial ball-and-socket joint between the head of the 
femur and the acetabulum of the hip bone ({link]). The hip carries the 
weight of the body and thus requires strength and stability during standing 
and walking. For these reasons, its range of motion is more limited than at 
the shoulder joint. 


The acetabulum is the socket portion of the hip joint. This space is deep and 
has a large articulation area for the femoral head, thus giving stability and 
weight bearing ability to the joint. The acetabulum is further deepened by 
the acetabular labrum, a fibrocartilage lip attached to the outer margin of 
the acetabulum. The surrounding articular capsule is strong, with several 


thickened areas forming intrinsic ligaments. These ligaments arise from the 
hip bone, at the margins of the acetabulum, and attach to the femur at the 
base of the neck. The ligaments are the iliofemoral ligament, pubofemoral 
ligament, and ischiofemoral ligament, all of which spiral around the head 
and neck of the femur. The ligaments are tightened by extension at the hip, 
thus pulling the head of the femur tightly into the acetabulum when in the 
upright, standing position. Very little additional extension of the thigh is 
permitted beyond this vertical position. These ligaments thus stabilize the 
hip joint and allow you to maintain an upright standing position with only 
minimal muscle contraction. Inside of the articular capsule, the ligament of 
the head of the femur (ligamentum teres) spans between the acetabulum 
and femoral head. This intracapsular ligament is normally slack and does 
not provide any significant joint support, but it does provide a pathway for 
an important artery that supplies the head of the femur. 


The hip is prone to osteoarthritis, and thus was the first joint for which a 
replacement prosthesis was developed. A common injury in elderly 
individuals, particularly those with weakened bones due to osteoporosis, is 
a “broken hip,” which is actually a fracture of the femoral neck. This may 
result from a fall, or it may cause the fall. This can happen as one lower 
limb is taking a step and all of the body weight is placed on the other limb, 
causing the femoral neck to break and producing a fall. Any accompanying 
disruption of the blood supply to the femoral neck or head can lead to 
necrosis of these areas, resulting in bone and cartilage death. Femoral 
fractures usually require surgical treatment, after which the patient will 
need mobility assistance for a prolonged period, either from family 
members or in a long-term care facility. Consequentially, the associated 
health care costs of “broken hips” are substantial. In addition, hip fractures 
are associated with increased rates of morbidity (incidences of disease) and 
mortality (death). Surgery for a hip fracture followed by prolonged bed rest 
may lead to life-threatening complications, including pneumonia, infection 
of pressure ulcers (bedsores), and thrombophlebitis (deep vein thrombosis; 
blood clot formation) that can result in a pulmonary embolism (blood clot 
within the lung). 

Hip Joint 


Articular cartilage Coxal (hip) bone 


Acetabular labrum 


Ligament of the 
head of the femur 


Synovial cavity 


Articular capsule 


(a) Frontal section through the right hip joint 


Anterior inferior 


A ) lliofemoral 
iliac spine ligament 
Greater 

tronchanter » “~% = \Os 
Pubofemoral . 

ligament 


(b) Anterior view of right hip joint, capsule in place 


Ischium 
v lliofemoral 
/ ean ligament 


_ Greater 


trochanter 
of femur 


Ischiofemoral 
ligament 


(c) Posterior view of right hip joint, capsule in place 


(a) The ball-and-socket joint of the 
hip is a multiaxial joint that provides 
both stability and a wide range of 
motion. (b—c) When standing, the 
supporting ligaments are tight, 
pulling the head of the femur into the 
acetabulum. 


. Loy 
wees Openstax COLLEGE 
= “ 

a= 

[=] fu 


Watch this video for a tutorial on the anatomy of the hip joint. What is a 
possible consequence following a fracture of the femoral neck within the 
capsule of the hip joint? 


Note: 


al: [=] 
7 
r. 

— 

meee OPENStAX COLLEGE 


1 


bese 


Watch this video to learn more about the anatomy of the hip joint, 
including bones, joints, muscles, nerves, and blood vessels. Where is the 
articular cartilage thickest within the hip joint? 


Knee Joint 


The knee joint is the largest joint of the body ([link]). It actually consists of 
three articulations. The femoropatellar joint is found between the patella 
and the distal femur. The medial tibiofemoral joint and lateral 
tibiofemoral joint are located between the medial and lateral condyles of 
the femur and the medial and lateral condyles of the tibia. All of these 
articulations are enclosed within a single articular capsule. The knee 
functions as a hinge joint, allowing flexion and extension of the leg. This 


action is generated by both rolling and gliding motions of the femur on the 
tibia. In addition, some rotation of the leg is available when the knee is 
flexed, but not when extended. The knee is well constructed for weight 
bearing in its extended position, but is vulnerable to injuries associated with 
hyperextension, twisting, or blows to the medial or lateral side of the joint, 
particularly while weight bearing. 


At the femoropatellar joint, the patella slides vertically within a groove on 
the distal femur. The patella is a sesamoid bone incorporated into the tendon 
of the quadriceps femoris muscle, the large muscle of the anterior thigh. 
The patella serves to protect the quadriceps tendon from friction against the 
distal femur. Continuing from the patella to the anterior tibia just below the 
knee is the patellar ligament. Acting via the patella and patellar ligament, 
the quadriceps femoris is a powerful muscle that acts to extend the leg at 
the knee. It also serves as a “dynamic ligament” to provide very important 
support and stabilization for the knee joint. 


The medial and lateral tibiofemoral joints are the articulations between the 
rounded condyles of the femur and the relatively flat condyles of the tibia. 
During flexion and extension motions, the condyles of the femur both roll 
and glide over the surfaces of the tibia. The rolling action produces flexion 
or extension, while the gliding action serves to maintain the femoral 
condyles centered over the tibial condyles, thus ensuring maximal bony, 
weight-bearing support for the femur in all knee positions. As the knee 
comes into full extension, the femur undergoes a slight medial rotation in 
relation to tibia. The rotation results because the lateral condyle of the 
femur is slightly smaller than the medial condyle. Thus, the lateral condyle 
finishes its rolling motion first, followed by the medial condyle. The 
resulting small medial rotation of the femur serves to “lock” the knee into 
its fully extended and most stable position. Flexion of the knee is initiated 
by a slight lateral rotation of the femur on the tibia, which “unlocks” the 
knee. This lateral rotation motion is produced by the popliteus muscle of the 
posterior leg. 


Located between the articulating surfaces of the femur and tibia are two 
articular discs, the medial meniscus and lateral meniscus (see [link]b). 
Each is a C-shaped fibrocartilage structure that is thin along its inside 


margin and thick along the outer margin. They are attached to their tibial 
condyles, but do not attach to the femur. While both menisci are free to 
move during knee motions, the medial meniscus shows less movement 
because it is anchored at its outer margin to the articular capsule and tibial 
collateral ligament. The menisci provide padding between the bones and 
help to fill the gap between the round femoral condyles and flattened tibial 
condyles. Some areas of each meniscus lack an arterial blood supply and 
thus these areas heal poorly if damaged. 


The knee joint has multiple ligaments that provide support, particularly in 
the extended position (see [link]c). Outside of the articular capsule, located 
at the sides of the knee, are two extrinsic ligaments. The fibular collateral 
ligament (lateral collateral ligament) is on the lateral side and spans from 
the lateral epicondyle of the femur to the head of the fibula. The tibial 
collateral ligament (medial collateral ligament) of the medial knee runs 
from the medial epicondyle of the femur to the medial tibia. As it crosses 
the knee, the tibial collateral ligament is firmly attached on its deep side to 
the articular capsule and to the medial meniscus, an important factor when 
considering knee injuries. In the fully extended knee position, both 
collateral ligaments are taut (tight), thus serving to stabilize and support the 
extended knee and preventing side-to-side or rotational motions between 
the femur and tibia. 


The articular capsule of the posterior knee is thickened by intrinsic 
ligaments that help to resist knee hyperextension. Inside the knee are two 
intracapsular ligaments, the anterior cruciate ligament and posterior 
cruciate ligament. These ligaments are anchored inferiorly to the tibia at 
the intercondylar eminence, the roughened area between the tibial condyles. 
The cruciate ligaments are named for whether they are attached anteriorly 
or posteriorly to this tibial region. Each ligament runs diagonally upward to 
attach to the inner aspect of a femoral condyle. The cruciate ligaments are 
named for the X-shape formed as they pass each other (cruciate means 
“cross”). The posterior cruciate ligament is the stronger ligament. It serves 
to support the knee when it is flexed and weight bearing, as when walking 
downhill. In this position, the posterior cruciate ligament prevents the femur 
from sliding anteriorly off the top of the tibia. The anterior cruciate 


ligament becomes tight when the knee is extended, and thus resists 


hyperextension. 
Knee Joint 


Femur Tendon of 


quadriceps femoris 


Articular 


Suprapatellar bursa 
capsule 


Patella 
Prepatellar bursa 


Posterior 
cruciate 
ligament Synovial cavity 

; Lateral meniscus 
meniscus 
Infrapatellar 


Anterior fat pad 


cruciate 
ligament 


Tibia 


Infrapatellar 
bursa 


Patellar ligament 


(a) Sagittal section through the right knee joint 


Quadriceps 
femoris muscle 


Tendon of —————————_ 
quadriceps 
femoris muscle 


a 


Medial patellar 


Lateral rosie |e Ay LX retinaculum 


retinaculum 
Z Tibial collateral 


Fibular collateral ligament 
ligament 

W! Patellar ligament 
Fibula = Tibia 


(c) Anterior view of right knee 


Anterior cruciate : 
ligament prienoe 
—_—— Articular 

cartilage on 
lateral tibial 


condyle 


Articular cartilage yo 
on medial F 


tibial condyle c 


Medial meniscus <<) Sf 


— Lateral 
meniscus 


\ <a 


Posterior cruciate 
ligament 


(b) Superior view of the right tibia in the knee joint, showing 
the menisci and cruciate ligaments 


(a) The knee joint is the largest joint of the body. (b)-(c) It is supported 
by the tibial and fibular collateral ligaments located on the sides of the 
knee outside of the articular capsule, and the anterior and posterior 
cruciate ligaments found inside the capsule. The medial and lateral 
menisci provide padding and support between the femoral condyles 
and tibial condyles. 


Note: 


Dees 0 
Per 


— 
mess" Openstax COLLEGE 


Watch this video to learn more about the flexion and extension of the knee, 
as the femur both rolls and glides on the tibia to maintain stable contact 
between the bones in all knee positions. The patella glides along a groove 
on the anterior side of the distal femur. The collateral ligaments on the 
sides of the knee become tight in the fully extended position to help 
stabilize the knee. The posterior cruciate ligament supports the knee when 
flexed and the anterior cruciate ligament becomes tight when the knee 
comes into full extension to resist hyperextension. What are the ligaments 
that support the knee joint? 


Note: 

ee 

_ openstax COLLEGE 

Oe 
Watch this video to learn more about the anatomy of the knee joint, 
including bones, joints, muscles, nerves, and blood vessels. Which 
ligament of the knee keeps the tibia from sliding too far forward in relation 


to the femur and which ligament keeps the tibia from sliding too far 
backward? 


Note: 
Disorders of the... 
Joints 


Injuries to the knee are common. Since this joint is primarily supported by 
muscles and ligaments, injuries to any of these structures will result in pain 
or knee instability. Injury to the posterior cruciate ligament occurs when 
the knee is flexed and the tibia is driven posteriorly, such as falling and 
landing on the tibial tuberosity or hitting the tibia on the dashboard when 
not wearing a seatbelt during an automobile accident. More commonly, 
injuries occur when forces are applied to the extended knee, particularly 
when the foot is planted and unable to move. Anterior cruciate ligament 
injuries can result with a forceful blow to the anterior knee, producing 
hyperextension, or when a runner makes a quick change of direction that 
produces both twisting and hyperextension of the knee. 

A worse combination of injuries can occur with a hit to the lateral side of 
the extended knee ([link]). A moderate blow to the lateral knee will cause 
the medial side of the joint to open, resulting in stretching or damage to the 
tibial collateral ligament. Because the medial meniscus is attached to the 
tibial collateral ligament, a stronger blow can tear the ligament and also 
damage the medial meniscus. This is one reason that the medial meniscus 
is 20 times more likely to be injured than the lateral meniscus. A powerful 
blow to the lateral knee produces a “terrible triad” injury, in which there is 
a sequential injury to the tibial collateral ligament, medial meniscus, and 
anterior cruciate ligament. 

Arthroscopic surgery has greatly improved the surgical treatment of knee 
injuries and reduced subsequent recovery times. This procedure involves a 
small incision and the insertion into the joint of an arthroscope, a pencil- 
thin instrument that allows for visualization of the joint interior. Small 
surgical instruments are also inserted via additional incisions. These tools 
allow a surgeon to remove or repair a torn meniscus or to reconstruct a 
ruptured cruciate ligament. The current method for anterior cruciate 
ligament replacement involves using a portion of the patellar ligament. 
Holes are drilled into the cruciate ligament attachment points on the tibia 
and femur, and the patellar ligament graft, with small areas of attached 
bone still intact at each end, is inserted into these holes. The bone-to-bone 
sites at each end of the graft heal rapidly and strongly, thus enabling a rapid 
recovery. 

Knee Injury 


Lateral Medial 


Torn tibial (medial) 


Direction of force collateral ligament 


Torn anterior Medial meniscus 
cruciate ligament 


Anterior view 


A strong blow to the lateral side of the extended 
knee will cause three injuries, in sequence: tearing 
of the tibial collateral ligament, damage to the 
medial meniscus, and rupture of the anterior 
cruciate ligament. 


Note: 
[=] [=] 
=h 
-—¥ openstax COLLEGE 
mina H 
[lr 


Watch this video to learn more about different knee injuries and diagnostic 
testing of the knee. What are the most common causes of anterior cruciate 


ligament injury? 


Ankle and Foot Joints 


The ankle is formed by the talocrural joint ((link]). It consists of the 
articulations between the talus bone of the foot and the distal ends of the 
tibia and fibula of the leg (crural = “leg”). The superior aspect of the talus 
bone is square-shaped and has three areas of articulation. The top of the 
talus articulates with the inferior tibia. This is the portion of the ankle joint 
that carries the body weight between the leg and foot. The sides of the talus 
are firmly held in position by the articulations with the medial malleolus of 
the tibia and the lateral malleolus of the fibula, which prevent any side-to- 
side motion of the talus. The ankle is thus a uniaxial hinge joint that allows 
only for dorsiflexion and plantar flexion of the foot. 


Additional joints between the tarsal bones of the posterior foot allow for the 
movements of foot inversion and eversion. Most important for these 
movements is the subtalar joint, located between the talus and calcaneus 
bones. The joints between the talus and navicular bones and the calcaneus 
and cuboid bones are also important contributors to these movements. All 
of the joints between tarsal bones are plane joints. Together, the small 
motions that take place at these joints all contribute to the production of 
inversion and eversion foot motions. 


Like the hinge joints of the elbow and knee, the talocrural joint of the ankle 
is supported by several strong ligaments located on the sides of the joint. 
These ligaments extend from the medial malleolus of the tibia or lateral 
malleolus of the fibula and anchor to the talus and calcaneus bones. Since 
they are located on the sides of the ankle joint, they allow for dorsiflexion 
and plantar flexion of the foot. They also prevent abnormal side-to-side and 
twisting movements of the talus and calcaneus bones during eversion and 
inversion of the foot. On the medial side is the broad deltoid ligament. The 
deltoid ligament supports the ankle joint and also resists excessive eversion 
of the foot. The lateral side of the ankle has several smaller ligaments. 
These include the anterior talofibular ligament and the posterior 
talofibular ligament, both of which span between the talus bone and the 
lateral malleolus of the fibula, and the calcaneofibular ligament, located 
between the calcaneus bone and fibula. These ligaments support the ankle 
and also resist excess inversion of the foot. 


Ankle Joint 


Tibia 


Medial malleolus 


Deltoid ligament 


Medial view 


Fibula Tibia 


Posterior and anterior inferior 


tibiofibular ligaments 
Interosseous 


membrane Anterior talofibular ligament 


Calcaneofibular ligament Subtalar joint 


Lateral view 


The talocrural (ankle) joint is a uniaxial 
hinge joint that only allows for 
dorsiflexion or plantar flexion of the foot. 
Movements at the subtalar joint, between 
the talus and calcaneus bones, combined 
with motions at other intertarsal joints, 
enables eversion/inversion movements of 
the foot. Ligaments that unite the medial 
or lateral malleolus with the talus and 
calcaneus bones serve to support the 
talocrural joint and to resist excess 
eversion or inversion of the foot. 


[=]: Le] 


Watch this video for a tutorial on the anatomy of the ankle joint. What are 
the three ligaments found on the lateral side of the ankle joint? 


Note: 
we 
fo 
= openstax COLLEGE 
fe eg 
[ele 


Watch this video to learn more about the anatomy of the ankle joint, 
including bones, joints, muscles, nerves, and blood vessels. Which type of 
joint used in woodworking does the ankle joint resemble? 


Note: 

Disorders of the... 

Joints 

The ankle is the most frequently injured joint in the body, with the most 
common injury being an inversion ankle sprain. A sprain is the stretching 
or tearing of the supporting ligaments. Excess inversion causes the talus 
bone to tilt laterally, thus damaging the ligaments on the lateral side of the 
ankle. The anterior talofibular ligament is most commonly injured, 
followed by the calcaneofibular ligament. In severe inversion injuries, the 


forceful lateral movement of the talus not only ruptures the lateral ankle 
ligaments, but also fractures the distal fibula. 

Less common are eversion sprains of the ankle, which involve stretching of 
the deltoid ligament on the medial side of the ankle. Forcible eversion of 
the foot, for example, with an awkward landing from a jump or when a 
football player has a foot planted and is hit on the lateral ankle, can result 
in a Pott’s fracture and dislocation of the ankle joint. In this injury, the very 
strong deltoid ligament does not tear, but instead shears off the medial 
malleolus of the tibia. This frees the talus, which moves laterally and 
fractures the distal fibula. In extreme cases, the posterior margin of the 
tibia may also be sheared off. 

Above the ankle, the distal ends of the tibia and fibula are united by a 
strong syndesmosis formed by the interosseous membrane and ligaments at 
the distal tibiofibular joint. These connections prevent separation between 
the distal ends of the tibia and fibula and maintain the talus locked into 
position between the medial malleolus and lateral malleolus. Injuries that 
produce a lateral twisting of the leg on top of the planted foot can result in 
stretching or tearing of the tibiofibular ligaments, producing a syndesmotic 
ankle sprain or “high ankle sprain.” 

Most ankle sprains can be treated using the RICE technique: Rest, Ice, 
Compression, and Elevation. Reducing joint mobility using a brace or cast 
may be required for a period of time. More severe injuries involving 
ligament tears or bone fractures may require surgery. 


Note: 


— 
mess" Openstax COLLEGE 


Watch this video to learn more about the ligaments of the ankle joint, ankle 
sprains, and treatment. During an inversion ankle sprain injury, all three 


ligaments that resist excessive inversion of the foot may be injured. What 
is the sequence in which these three ligaments are injured? 


Chapter Review 


Although synovial joints share many common features, each joint of the 
body is specialized for certain movements and activities. The joints of the 
upper limb provide for large ranges of motion, which give the upper limb 
great mobility, thus enabling actions such as the throwing of a ball or typing 
on a keyboard. The joints of the lower limb are more robust, giving them 
greater strength and the stability needed to support the body weight during 
running, jumping, or kicking activities. 


The joints of the vertebral column include the symphysis joints formed by 
each intervertebral disc and the plane synovial joints between the superior 
and inferior articular processes of adjacent vertebrae. Each of these joints 
provide for limited motions, but these sum together to produce flexion, 
extension, lateral flexion, and rotation of the neck and body. The range of 
motions available in each region of the vertebral column varies, with all of 
these motions available in the cervical region. Only rotation is allowed in 
the thoracic region, while the lumbar region has considerable extension, 
flexion, and lateral flexion, but rotation is prevented. The atlanto-occipital 
joint allows for flexion and extension of the head, while the atlantoaxial 
joint is a pivot joint that provides for rotation of the head. 


The temporomandibular joint is the articulation between the condyle of the 
mandible and the mandibular fossa and articular tubercle of the skull 
temporal bone. An articular disc is located between the bony components of 
this joint. A combination of gliding and hinge motions of the mandibular 
condyle allows for elevation/depression, protraction/retraction, and side-to- 
side motions of the lower jaw. 


The glenohumeral (shoulder) joint is a multiaxial ball-and-socket joint that 
provides flexion/extension, abduction/adduction, circumduction, and 

medial/lateral rotation of the humerus. The head of the humerus articulates 
with the glenoid cavity of the scapula. The glenoid labrum extends around 


the margin of the glenoid cavity. Intrinsic ligaments, including the 
coracohumeral ligament and glenohumeral ligaments, provide some support 
for the shoulder joint. However, the primary support comes from muscles 
crossing the joint whose tendons form the rotator cuff. These muscle 
tendons are protected from friction against the scapula by the subacromial 
bursa and subscapular bursa. 


The elbow is a uniaxial hinge joint that allows for flexion/extension of the 
forearm. It includes the humeroulnar joint and the humeroradial joint. The 
medial elbow is supported by the ulnar collateral ligament and the radial 
collateral ligament supports the lateral side. These ligaments prevent side- 
to-side movements and resist hyperextension of the elbow. The proximal 
radioulnar joint is a pivot joint that allows for rotation of the radius during 
pronation/supination of the forearm. The annular ligament surrounds the 
head of the radius to hold it in place at this joint. 


The hip joint is a ball-and-socket joint whose motions are more restricted 
than at the shoulder to provide greater stability during weight bearing. The 
hip joint is the articulation between the head of the femur and the 
acetabulum of the hip bone. The acetabulum is deepened by the acetabular 
labrum. The iliofemoral, pubofemoral, and ischiofemoral ligaments 
strongly support the hip joint in the upright, standing position. The ligament 
of the head of the femur provides little support but carries an important 
artery that supplies the femur. 


The knee includes three articulations. The femoropatellar joint is between 
the patella and distal femur. The patella, a sesamoid bone incorporated into 
the tendon of the quadriceps femoris muscle of the anterior thigh, serves to 
protect this tendon from rubbing against the distal femur during knee 
movements. The medial and lateral tibiofemoral joints, between the 
condyles of the femur and condyles of the tibia, are modified hinge joints 
that allow for knee extension and flexion. During these movements, the 
condyles of the femur both roll and glide over the surface of the tibia. As 
the knee comes into full extension, a slight medial rotation of the femur 
serves to “lock” the knee into its most stable, weight-bearing position. The 
reverse motion, a small lateral rotation of the femur, is required to initiate 
knee flexion. When the knee is flexed, some rotation of the leg is available. 


Two extrinsic ligaments, the tibial collateral ligament on the medial side 
and the fibular collateral ligament on the lateral side, serve to resist 
hyperextension or rotation of the extended knee joint. Two intracapsular 
ligaments, the anterior cruciate ligament and posterior cruciate ligament, 
span between the tibia and the inner aspects of the femoral condyles. The 
anterior cruciate ligament resists hyperextension of the knee, while the 
posterior cruciate ligament prevents anterior sliding of the femur, thus 
supporting the knee when it is flexed and weight bearing. The medial and 
lateral menisci, located between the femoral and tibial condyles, are 
articular discs that provide padding and improve the fit between the bones. 


The talocrural joint forms the ankle. It consists of the articulation between 
the talus bone and the medial malleolus of the tibia, the distal end of the 
tibia, and the lateral malleolus of the fibula. This is a uniaxial hinge joint 
that allows only dorsiflexion and plantar flexion of the foot. Gliding 
motions at the subtalar and intertarsal joints of the foot allow for 
inversion/eversion of the foot. The ankle joint is supported on the medial 
side by the deltoid ligament, which prevents side-to-side motions of the 
talus at the talocrural joint and resists excessive eversion of the foot. The 
lateral ankle is supported by the anterior and posterior talofibular ligaments 
and the calcaneofibular ligament. These support the ankle joint and also 
resist excess inversion of the foot. An inversion ankle sprain, a common 
injury, will result in injury to one or more of these lateral ankle ligaments. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to learn about TMJ. Opening of the mouth requires 
the combination of two motions at the temporomandibular joint, an 
anterior gliding motion of the articular disc and mandible and the 
downward hinging of the mandible. What is the initial movement of 
the mandible during opening and how much mouth opening does this 
produce? 


Solution: 


The first motion is rotation (hinging) of the mandible, but this only 
produces about 20 mm (0.78 in) of mouth opening. 


Exercise: 
Problem: 


Watch this video for a tutorial on the anatomy of the shoulder joint. 
What movements are available at the shoulder joint? 


Solution: 


The shoulder joint is a ball-and-socket joint that allows for flexion- 
extension, abduction-adduction, medial rotation, lateral rotation, and 
circumduction of the humerus. 


Exercise: 
Problem: 
Watch this video to learn about the anatomy of the shoulder joint, 
including bones, joints, muscles, nerves, and blood vessels. What is the 


shape of the glenoid labrum in cross-section, and what is the 
importance of this shape? 


Solution: 


The glenoid labrum is wedge-shaped in cross-section. This is 
important because it creates an elevated rim around the glenoid cavity, 
which creates a deeper socket for the head of the humerus to fit into. 


Exercise: 


Problem: 


Watch this animation to learn more about the anatomy of the elbow 
joint. What structures provide the main stability for the elbow? 


Solution: 


The structures that stabilize the elbow include the coronoid process, 
the radial (lateral) collateral ligament, and the anterior portion of the 
ulnar (medial) collateral ligament. 


Exercise: 
Problem: 
Watch this video to learn more about the anatomy of the elbow joint, 


including bones, joints, muscles, nerves, and blood vessels. What are 
the functions of the articular cartilage? 


Solution: 


The articular cartilage functions to absorb shock and to provide an 
extremely smooth surface that makes movement between bones easy, 
without damaging the bones. 


Exercise: 
Problem: 
Watch this video for a tutorial on the anatomy of the hip joint. What is 


a possible consequence following a fracture of the femoral neck within 
the capsule of the hip joint? 


Solution: 


An intracapsular fracture of the neck of the femur can result in 
disruption of the arterial blood supply to the head of the femur, which 
may lead to avascular necrosis of the femoral head. 


Exercise: 
Problem: 
Watch this video to learn more about the anatomy of the hip joint, 


including bones, joints, muscles, nerves, and blood vessels. Where is 
the articular cartilage thickest within the hip joint? 


Solution: 


The articular cartilage is thickest in the upper and back part of the 
acetabulum, the socket portion of the hip joint. These regions receive 
most of the force from the head of the femur during walking and 
running. 


Exercise: 


Problem: 


Watch this video to learn more about the flexion and extension of the 
knee, as the femur both rolls and glides on the tibia to maintain stable 
contact between the bones in all knee positions. The patella glides 
along a groove on the anterior side of the distal femur. The collateral 
ligaments on the sides of the knee become tight in the fully extended 
position to help stabilize the knee. The posterior cruciate ligament 
supports the knee when flexed and the anterior cruciate ligament 
becomes tight when the knee comes into full extension to resist 
hyperextension. What are the ligaments that support the knee joint? 


Solution: 


There are five ligaments associated with the knee joint. The tibial 
collateral ligament is located on the medial side of the knee and the 
fibular collateral ligament is located on the lateral side. The anterior 
and posterior cruciate ligaments are located inside the knee joint. 


Exercise: 
Problem: 
Watch this video to learn more about the anatomy of the knee joint, 
including bones, joints, muscles, nerves, and blood vessels. Which 
ligament of the knee keeps the tibia from sliding too far forward in 


relation to the femur and which ligament keeps the tibia from sliding 
too far backward? 


Solution: 


The anterior cruciate ligament prevents the tibia from sliding too far 
forward in relation to the femur and the posterior cruciate ligament 


keeps the tibia from sliding too far backward. 
Exercise: 
Problem: 
Watch this video to learn more about different knee injuries and 


diagnostic testing of the knee. What are the most causes of anterior 
cruciate ligament injury? 


Solution: 


The anterior cruciate ligament (ACL) is most commonly injured when 
traumatic force is applied to the knee during a twisting motion or when 
side standing or landing from a jump. 


Exercise: 
Problem: 


Watch this video for a tutorial on the anatomy of the ankle joint. What 
are the three ligaments found on the lateral side of the ankle joint? 


Solution: 


The ligaments of the lateral ankle are the anterior and posterior 
talofibular ligaments and the calcaneofibular ligament. These 
ligaments support the ankle joint and resist excess inversion of the 
foot. 


Exercise: 


Problem: 


Watch this video to learn more about the anatomy of the ankle joint, 
including bones, joints, muscles, nerves, and blood vessels. The ankle 
joint resembles what type of joint used in woodworking? 


Solution: 


Because of the square shape of the ankle joint, it has been compared to 
a mortise-and-tendon type of joint. 


Exercise: 
Problem: 
Watch this video to learn about the ligaments of the ankle joint, ankle 
sprains, and treatment. During an inversion ankle sprain injury, all 
three ligaments that resist excessive inversion of the foot may be 


injured. What is the sequence in which these three ligaments are 
injured? 


Solution: 


An inversion ankle sprain may injure all three ligaments located on the 
lateral side of the ankle. The sequence of injury would be the anterior 
talofibular ligament first, followed by the calcaneofibular ligament 
second, and finally, the posterior talofibular ligament third. 


Review Questions 


Exercise: 


Problem: 
The primary support for the glenohumeral joint is provided by the 
a. coracohumeral ligament 
b. glenoid labrum 
c. rotator cuff muscles 
d. subacromial bursa 
Solution: 


C 


Exercise: 


Problem: The proximal radioulnar joint 


a. is supported by the annular ligament 

b. contains an articular disc that strongly unites the bones 

c. is supported by the ulnar collateral ligament 

d. is a hinge joint that allows for flexion/extension of the forearm 


Solution: 


A 


Exercise: 


Problem: Which statement is true concerning the knee joint? 


a. The lateral meniscus is an intrinsic ligament located on the lateral 
side of the knee joint. 

b. Hyperextension is resisted by the posterior cruciate ligament. 

c. The anterior cruciate ligament supports the knee when it is flexed 
and weight bearing. 

d. The medial meniscus is attached to the tibial collateral ligament. 


Solution: 


D 


Exercise: 


Problem:The ankle joint 


a. is also called the subtalar joint 

b. allows for gliding movements that produce inversion/eversion of 
the foot 

c. is a uniaxial hinge joint 


d. is supported by the tibial collateral ligament on the lateral side 


Solution: 


C 
Exercise: 


Problem: 


Which region of the vertebral column has the greatest range of motion 
for rotation? 


a. cervical 
b. thoracic 
c. lumbar 
d. sacral 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


Discuss the structures that contribute to support of the shoulder joint. 


Solution: 


The shoulder joint allows for a large range of motion. The primary 
support for the shoulder joint is provided by the four rotator cuff 
muscles. These muscles serve as “dynamic ligaments” and thus can 
modulate their strengths of contraction as needed to hold the head of 
the humerus in position at the glenoid fossa. Additional but weaker 


support comes from the coracohumeral ligament, an intrinsic ligament 
that supports the superior aspect of the shoulder joint, and the 
glenohumeral ligaments, which are intrinsic ligaments that support the 
anterior side of the joint. 


Exercise: 


Problem: 


Describe the sequence of injuries that may occur if the extended, 
weight-bearing knee receives a very strong blow to the lateral side of 
the knee. 


Solution: 


A strong blow to the lateral side of the extended knee will cause the 
medial side of the knee joint to open, resulting in a sequence of three 
injuries. First will be damage to the tibial collateral ligament. Since the 
medial meniscus is attached to the tibial collateral ligament, the 
meniscus is also injured. The third structure injured would be the 
anterior cruciate ligament. 


Glossary 


acetabular labrum 
lip of fibrocartilage that surrounds outer margin of the acetabulum on 
the hip bone 


annular ligament 
intrinsic ligament of the elbow articular capsule that surrounds and 
supports the head of the radius at the proximal radioulnar joint 


anterior cruciate ligament 
intracapsular ligament of the knee; extends from anterior, superior 
surface of the tibia to the inner aspect of the lateral condyle of the 
femur; resists hyperextension of knee 


anterior talofibular ligament 


intrinsic ligament located on the lateral side of the ankle joint, between 
talus bone and lateral malleolus of fibula; supports talus at the 
talocrural joint and resists excess inversion of the foot 


atlantoaxial joint 
series of three articulations between the atlas (C1) vertebra and the 
axis (C2) vertebra, consisting of the joints between the inferior 
articular processes of C1 and the superior articular processes of C2, 
and the articulation between the dens of C2 and the anterior arch of C1 


atlanto-occipital joint 
articulation between the occipital condyles of the skull and the 
superior articular processes of the atlas (C1 vertebra) 


calcaneofibular ligament 
intrinsic ligament located on the lateral side of the ankle joint, between 
the calcaneus bone and lateral malleolus of the fibula; supports the 
talus bone at the ankle joint and resists excess inversion of the foot 


coracohumeral ligament 
intrinsic ligament of the shoulder joint; runs from the coracoid process 
of the scapula to the anterior humerus 


deltoid ligament 
broad intrinsic ligament located on the medial side of the ankle joint; 
supports the talus at the talocrural joint and resists excess eversion of 
the foot 


elbow joint 
humeroulnar joint 


femoropatellar joint 
portion of the knee joint consisting of the articulation between the 
distal femur and the patella 


fibular collateral ligament 
extrinsic ligament of the knee joint that spans from the lateral 
epicondyle of the femur to the head of the fibula; resists 


hyperextension and rotation of the extended knee 


glenohumeral joint 
shoulder joint; articulation between the glenoid cavity of the scapula 
and head of the humerus; multiaxial ball-and-socket joint that allows 
for flexion/extension, abduction/adduction, circumduction, and 
medial/lateral rotation of the humerus 


glenohumeral ligament 
one of the three intrinsic ligaments of the shoulder joint that strengthen 
the anterior articular capsule 


glenoid labrum 
lip of fibrocartilage located around the outside margin of the glenoid 
cavity of the scapula 


humeroradial joint 
articulation between the capitulum of the humerus and head of the 
radius 


humeroulnar joint 
articulation between the trochlea of humerus and the trochlear notch of 
the ulna; uniaxial hinge joint that allows for flexion/extension of the 
forearm 


iliofemoral ligament 
intrinsic ligament spanning from the ilium of the hip bone to the 
femur, on the superior-anterior aspect of the hip joint 


ischiofemoral ligament 
intrinsic ligament spanning from the ischium of the hip bone to the 
femur, on the posterior aspect of the hip joint 


lateral meniscus 
C-shaped fibrocartilage articular disc located at the knee, between the 
lateral condyle of the femur and the lateral condyle of the tibia 


lateral tibiofemoral joint 


portion of the knee consisting of the articulation between the lateral 
condyle of the tibia and the lateral condyle of the femur; allows for 
flexion/extension at the knee 


ligament of the head of the femur 
intracapsular ligament that runs from the acetabulum of the hip bone to 
the head of the femur 


medial meniscus 
C-shaped fibrocartilage articular disc located at the knee, between the 
medial condyle of the femur and medial condyle of the tibia 


medial tibiofemoral joint 
portion of the knee consisting of the articulation between the medial 
condyle of the tibia and the medial condyle of the femur; allows for 
flexion/extension at the knee 


patellar ligament 
ligament spanning from the patella to the anterior tibia; serves as the 
final attachment for the quadriceps femoris muscle 


posterior cruciate ligament 
intracapsular ligament of the knee; extends from the posterior, superior 
surface of the tibia to the inner aspect of the medial condyle of the 
femur; prevents anterior displacement of the femur when the knee is 
flexed and weight bearing 


posterior talofibular ligament 
intrinsic ligament located on the lateral side of the ankle joint, between 
the talus bone and lateral malleolus of the fibula; supports the talus at 
the talocrural joint and resists excess inversion of the foot 


pubofemoral ligament 
intrinsic ligament spanning from the pubis of the hip bone to the 
femur, on the anterior-inferior aspect of the hip joint 


radial collateral ligament 


intrinsic ligament on the lateral side of the elbow joint; runs from the 
lateral epicondyle of humerus to merge with the annular ligament 


rotator cuff 
strong connective tissue structure formed by the fusion of four rotator 
cuff muscle tendons to the articular capsule of the shoulder joint; 
surrounds and supports superior, anterior, lateral, and posterior sides of 
the humeral head 


subacromial bursa 
bursa that protects the supraspinatus muscle tendon and superior end 
of the humerus from rubbing against the acromion of the scapula 


subscapular bursa 
bursa that prevents rubbing of the subscapularis muscle tendon against 
the scapula 


subtalar joint 
articulation between the talus and calcaneus bones of the foot; allows 
motions that contribute to inversion/eversion of the foot 


talocrural joint 
ankle joint; articulation between the talus bone of the foot and medial 
malleolus of the tibia, distal tibia, and lateral malleolus of the fibula; a 
uniaxial hinge joint that allows only for dorsiflexion and plantar 
flexion of the foot 


temporomandibular joint (TMJ) 
articulation between the condyle of the mandible and the mandibular 
fossa and articular tubercle of the temporal bone of the skull; allows 
for depression/elevation (opening/closing of mouth), 
protraction/retraction, and side-to-side motions of the mandible 


tibial collateral ligament 
extrinsic ligament of knee joint that spans from the medial epicondyle 
of the femur to the medial tibia; resists hyperextension and rotation of 
extended knee 


ulnar collateral ligament 
intrinsic ligament on the medial side of the elbow joint; spans from the 
medial epicondyle of the humerus to the medial ulna 


zygapophysial joints 
facet joints; plane joints between the superior and inferior articular 
processes of adjacent vertebrae that provide for only limited motions 
between the vertebrae 


Overview of Muscle Tissues 
By the end of this section, you will be able to: 


e Describe the different types of muscle 
e Explain contractibility and extensibility 


Muscle is one of the four primary tissue types of the body, and the body 
contains three types of muscle tissue: skeletal muscle, cardiac muscle, and 
smooth muscle ([link]). All three muscle tissues have some properties in 
common; they all exhibit a quality called excitability as their plasma 
membranes can change their electrical states (from polarized to 
depolarized) and send an electrical wave called an action potential along the 
entire length of the membrane. While the nervous system can influence the 
excitability of cardiac and smooth muscle to some degree, skeletal muscle 
completely depends on signaling from the nervous system to work properly. 
On the other hand, both cardiac muscle and smooth muscle can respond to 
other stimuli, such as hormones and local stimuli. 

The Three Types of Muscle Tissue 


The body contains three types of 
muscle tissue: (a) skeletal muscle, 
(b) smooth muscle, and (c) cardiac 

muscle. From top, LM x 1600, 
LM x 1600, LM x 1600. 
(Micrographs provided by the 
Regents of University of Michigan 
Medical School © 2012) 


The muscles all begin the actual process of contracting (shortening) when a 
protein called actin is pulled by a protein called myosin. This occurs in 


striated muscle (skeletal and cardiac) after specific binding sites on the actin 
have been exposed in response to the interaction between calcium ions 
(Ca**) and proteins (troponin and tropomyosin) that “shield” the actin- 
binding sites. Ca** also is required for the contraction of smooth muscle, 
although its role is different: here Ca** activates enzymes, which in turn 
activate myosin heads. All muscles require adenosine triphosphate (ATP) to 
continue the process of contracting, and they all relax when the Ca** is 
removed and the actin-binding sites are re-shielded. 


A muscle can return to its original length when relaxed due to a quality of 
muscle tissue called elasticity. It can recoil back to its original length due to 
elastic fibers. Muscle tissue also has the quality of extensibility; it can 
stretch or extend. Contractility allows muscle tissue to pull on its 
attachment points and shorten with force. 


Differences among the three muscle types include the microscopic 
organization of their contractile proteins—actin and myosin. The actin and 
myosin proteins are arranged very regularly in the cytoplasm of individual 
muscle cells (referred to as fibers) in both skeletal muscle and cardiac 
muscle, which creates a pattern, or stripes, called striations. The striations 
are visible with a light microscope under high magnification (see [link]). 
Skeletal muscle fibers are multinucleated structures that compose the 
skeletal muscle. Cardiac muscle fibers each have one to two nuclei and are 
physically and electrically connected to each other so that the entire heart 
contracts as one unit (called a syncytium). 


Because the actin and myosin are not arranged in such regular fashion in 
smooth muscle, the cytoplasm of a smooth muscle fiber (which has only a 
single nucleus) has a uniform, nonstriated appearance (resulting in the name 
smooth muscle). However, the less organized appearance of smooth muscle 
should not be interpreted as less efficient. Smooth muscle in the walls of 
arteries is a critical component that regulates blood pressure necessary to 
push blood through the circulatory system; and smooth muscle in the skin, 
visceral organs, and internal passageways is essential for moving all 
materials through the body. 


Chapter Review 


Muscle is the tissue in animals that allows for active movement of the body 
or materials within the body. There are three types of muscle tissue: skeletal 
muscle, cardiac muscle, and smooth muscle. Most of the body’s skeletal 
muscle produces movement by acting on the skeleton. Cardiac muscle is 
found in the wall of the heart and pumps blood through the circulatory 
system. 


Smooth muscle is found in the skin, where it is associated with hair 
follicles; it also is found in the walls of internal organs, blood vessels, and 
internal passageways, where it assists in moving materials. 


Review Questions 


Exercise: 


Problem: 
Muscle that has a striped appearance is described as being 


a. elastic 

b. nonstriated 
c. excitable 
d. striated 


Solution: 


D 
Exercise: 


Problem: 
Which element is important in directly triggering contraction? 


a. sodium (Na*) 
b. calcium (Ca™) 
c. potassium (K*) 
d. chloride (CI) 


Solution: 


B 
Exercise: 


Problem: 


Which of the following properties is not common to all three muscle 
tissues? 


a. excitability 

b. the need for ATP 

c. at rest, uses shielding proteins to cover actin-binding sites 
d. elasticity 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: Why is elasticity an important quality of muscle tissue? 
Solution: 
It allows muscle to return to its original length during relaxation after 
contraction. 

Glossary 


cardiac muscle 


striated muscle found in the heart; joined to one another at intercalated 
discs and under the regulation of pacemaker cells, which contract as 
one unit to pump blood through the circulatory system. Cardiac muscle 
is under involuntary control. 


contractility 
ability to shorten (contract) forcibly 


elasticity 
ability to stretch and rebound 


excitability 
ability to undergo neural stimulation 


extensibility 
ability to lengthen (extend) 


skeletal muscle 
striated, multinucleated muscle that requires signaling from the 
nervous system to trigger contraction; most skeletal muscles are 
referred to as voluntary muscles that move bones and produce 
movement 


smooth muscle 
nonstriated, mononucleated muscle in the skin that is associated with 
hair follicles; assists in moving materials in the walls of internal 
organs, blood vessels, and internal passageways 


Skeletal Muscle 
By the end of this section, you will be able to: 


¢ Describe the layers of connective tissues packaging skeletal muscle 
e Explain how muscles work with tendons to move the body 

e Identify areas of the skeletal muscle fibers 

e Describe excitation-contraction coupling 


The best-known feature of skeletal muscle is its ability to contract and 
cause movement. Skeletal muscles act not only to produce movement but 
also to stop movement, such as resisting gravity to maintain posture. Small, 
constant adjustments of the skeletal muscles are needed to hold a body 
upright or balanced in any position. Muscles also prevent excess movement 
of the bones and joints, maintaining skeletal stability and preventing 
skeletal structure damage or deformation. Joints can become misaligned or 
dislocated entirely by pulling on the associated bones; muscles work to 
keep joints stable. Skeletal muscles are located throughout the body at the 
openings of internal tracts to control the movement of various substances. 
These muscles allow functions, such as swallowing, urination, and 
defecation, to be under voluntary control. Skeletal muscles also protect 
internal organs (particularly abdominal and pelvic organs) by acting as an 
external barrier or shield to external trauma and by supporting the weight of 
the organs. 


Skeletal muscles contribute to the maintenance of homeostasis in the body 
by generating heat. Muscle contraction requires energy, and when ATP is 
broken down, heat is produced. This heat is very noticeable during exercise, 
when sustained muscle movement causes body temperature to rise, and in 
cases of extreme cold, when shivering produces random skeletal muscle 
contractions to generate heat. 


Each skeletal muscle is an organ that consists of various integrated tissues. 
These tissues include the skeletal muscle fibers, blood vessels, nerve fibers, 
and connective tissue. Each skeletal muscle has three layers of connective 
tissue (called “mysia”) that enclose it and provide structure to the muscle as 
a whole, and also compartmentalize the muscle fibers within the muscle 
({link]). Each muscle is wrapped in a sheath of dense, irregular connective 
tissue called the epimysium, which allows a muscle to contract and move 


powerfully while maintaining its structural integrity. The epimysium also 
separates muscle from other tissues and organs in the area, allowing the 
muscle to move independently. 


The Three Connective Tissue Layers 
Skeletal muscle Epimysium Muscle fascicles 


Perimysium 
Endomysium 
Muscle fibers 


Muscle fascicle 


Muscle fiber 


Sarcolemma 


Bundles of muscle fibers, called fascicles, are 
covered by the perimysium. Muscle fibers are 
covered by the endomysium. 


Inside each skeletal muscle, muscle fibers are organized into individual 
bundles, each called a fascicle, by a middle layer of connective tissue called 
the perimysium. This fascicular organization is common in muscles of the 
limbs; it allows the nervous system to trigger a specific movement of a 
muscle by activating a subset of muscle fibers within a bundle, or fascicle 
of the muscle. Inside each fascicle, each muscle fiber is encased in a thin 


connective tissue layer of collagen and reticular fibers called the 
endomysium. The endomysium contains the extracellular fluid and 
nutrients to support the muscle fiber. These nutrients are supplied via blood 
to the muscle tissue. 


In skeletal muscles that work with tendons to pull on bones, the collagen in 
the three tissue layers (the mysia) intertwines with the collagen of a tendon. 
At the other end of the tendon, it fuses with the periosteum coating the 
bone. The tension created by contraction of the muscle fibers is then 
transferred though the mysia, to the tendon, and then to the periosteum to 
pull on the bone for movement of the skeleton. In other places, the mysia 
may fuse with a broad, tendon-like sheet called an aponeurosis, or to 
fascia, the connective tissue between skin and bones. The broad sheet of 
connective tissue in the lower back that the latissimus dorsi muscles (the 
“lats”) fuse into is an example of an aponeurosis. 


Every skeletal muscle is also richly supplied by blood vessels for 
nourishment, oxygen delivery, and waste removal. In addition, every 
muscle fiber in a skeletal muscle is supplied by the axon branch of a 
somatic motor neuron, which signals the fiber to contract. Unlike cardiac 
and smooth muscle, the only way to functionally contract a skeletal muscle 
is through signaling from the nervous system. 


Skeletal Muscle Fibers 


Because skeletal muscle cells are long and cylindrical, they are commonly 
referred to as muscle fibers. Skeletal muscle fibers can be quite large for 
human cells, with diameters up to 100 ym and lengths up to 30 cm (11.8 in) 
in the Sartorius of the upper leg. During early development, embryonic 
myoblasts, each with its own nucleus, fuse with up to hundreds of other 
myoblasts to form the multinucleated skeletal muscle fibers. Multiple nuclei 
mean multiple copies of genes, permitting the production of the large 
amounts of proteins and enzymes needed for muscle contraction. 


Some other terminology associated with muscle fibers is rooted in the 
Greek sarco, which means “flesh.” The plasma membrane of muscle fibers 
is called the sarcolemma, the cytoplasm is referred to as sarcoplasm, and 


the specialized smooth endoplasmic reticulum, which stores, releases, and 
retrieves calcium ions (Ca™*) is called the sarcoplasmic reticulum (SR) 
({link]). As will soon be described, the functional unit of a skeletal muscle 
fiber is the sarcomere, a highly organized arrangement of the contractile 
myofilaments actin (thin filament) and myosin (thick filament), along with 
other support proteins. 

Muscle Fiber 


Nucleus Muscle fiber a 
——— eg _C 


Mitochondrion 
Sarcolemma 


Light | 
band 


Dark A band 


Sarcoplasmic Sarcomere 
reticulum 


Thin (actin) Thick (myosin) 
filament Z disc H zone Z disc filament 


| band A band | band M line 


A skeletal muscle fiber is surrounded by a 
plasma membrane called the sarcolemma, 
which contains sarcoplasm, the cytoplasm of 
muscle cells. A muscle fiber is composed of 
many fibrils, which give the cell its striated 
appearance. 


The Sarcomere 


The striated appearance of skeletal muscle fibers is due to the arrangement 
of the myofilaments of actin and myosin in sequential order from one end 
of the muscle fiber to the other. Each packet of these microfilaments and 


their regulatory proteins, troponin and tropomyosin (along with other 
proteins) is called a sarcomere. 


Note: 


- 
openstax COLLEGE” 


Watch this video to learn more about macro- and microstructures of 
skeletal muscles. (a) What are the names of the “junction points” between 
sarcomeres? (b) What are the names of the “subunits” within the 
myofibrils that run the length of skeletal muscle fibers? (c) What is the 
“double strand of pearls” described in the video? (d) What gives a skeletal 
muscle fiber its striated appearance? 


The sarcomere is the functional unit of the muscle fiber. The sarcomere 
itself is bundled within the myofibril that runs the entire length of the 
muscle fiber and attaches to the sarcolemma at its end. As myofibrils 
contract, the entire muscle cell contracts. Because myofibrils are only 
approximately 1.2 ym in diameter, hundreds to thousands (each with 
thousands of sarcomeres) can be found inside one muscle fiber. Each 
sarcomere is approximately 2 pm in length with a three-dimensional 
cylinder-like arrangement and is bordered by structures called Z-discs (also 
called Z-lines, because pictures are two-dimensional), to which the actin 
myofilaments are anchored ({link]). Because the actin and its troponin- 
tropomyosin complex (projecting from the Z-discs toward the center of the 
sarcomere) form strands that are thinner than the myosin, it is called the 
thin filament of the sarcomere. Likewise, because the myosin strands and 
their multiple heads (projecting from the center of the sarcomere, toward 
but not all to way to, the Z-discs) have more mass and are thicker, they are 
called the thick filament of the sarcomere. 


The Sarcomere 


Sarcomere 


H zone 


Lighter! band Darker A band Lighter | band 


Portion of a Portion of a 
thick filament thin filament 


| Troponin Actin — Tropomyosin 


mi = 4 — ax 


Actin-binding sites 


Binding site Actin subunits 
for myosin 


ATP-binding site 
Tail Heads 


Myosin molecule Flexible hinge region 


The sarcomere, the region from one Z-line 
to the next Z-line, is the functional unit of a 
skeletal muscle fiber. 


The Neuromuscular Junction 


Another specialization of the skeletal muscle is the site where a motor 
neuron’s terminal meets the muscle fiber—called the neuromuscular 
junction (NMJ). This is where the muscle fiber first responds to signaling 
by the motor neuron. Every skeletal muscle fiber in every skeletal muscle is 
innervated by a motor neuron at the NMJ. Excitation signals from the 
neuron are the only way to functionally activate the fiber to contract. 


Note: 


— : 
mess Openstax COLLEGE 


parr. ir nl 
Every skeletal muscle fiber is supplied by a motor neuron at the NMJ. 
Watch this video to learn more about what happens at the NMJ. (a) What is 
the definition of a motor unit? (b) What is the structural and functional 
difference between a large motor unit and a small motor unit? (c) Can you 


give an example of each? (d) Why is the neurotransmitter acetylcholine 
degraded after binding to its receptor? 


Excitation-Contraction Coupling 


All living cells have membrane potentials, or electrical gradients across 
their membranes. The inside of the membrane is usually around -60 to -90 
mV, relative to the outside. This is referred to as a cell’s membrane 
potential. Neurons and muscle cells can use their membrane potentials to 
generate electrical signals. They do this by controlling the movement of 
charged particles, called ions, across their membranes to create electrical 
currents. This is achieved by opening and closing specialized proteins in the 
membrane called ion channels. Although the currents generated by ions 
moving through these channel proteins are very small, they form the basis 
of both neural signaling and muscle contraction. 


Both neurons and skeletal muscle cells are electrically excitable, meaning 
that they are able to generate action potentials. An action potential is a 
special type of electrical signal that can travel along a cell membrane as a 
wave. This allows a signal to be transmitted quickly and faithfully over long 
distances. 


Although the term excitation-contraction coupling confuses or scares 
some students, it comes down to this: for a skeletal muscle fiber to contract, 
its membrane must first be “excited”—in other words, it must be stimulated 


to fire an action potential. The muscle fiber action potential, which sweeps 
along the sarcolemma as a wave, is “coupled” to the actual contraction 
through the release of calcium ions (Ca**) from the SR. Once released, the 
Ca** interacts with the shielding proteins, forcing them to move aside so 
that the actin-binding sites are available for attachment by myosin heads. 
The myosin then pulls the actin filaments toward the center, shortening the 
muscle fiber. 


In skeletal muscle, this sequence begins with signals from the somatic 
motor division of the nervous system. In other words, the “excitation” step 
in skeletal muscles is always triggered by signaling from the nervous 
system ([link]). 

Motor End-Plate and Innervation 


Myelin sheath surrounding 
axon of motor neuron 


Axon terminal 


Synaptic end bulb at the 


Sarcolemma neuromuscular junction 


Myofibril of 
muscle fiber 


Sarcoplasm 


Synaptic end bulb 


Synaptic vesicle 


containing ACh Nerve impulse 


(action potential) 


Sarcolemma 


Synaptic 
cleft 


Motor end- 
plate 


ACh 
Synaptic 
vesicle 
Synaptic 
vesicle 
releases 
ACh by e° : 
exocytosis __ 9 = Synaptic cleft 


ACh receptor 


Motor 


Binding of end-plate 


ACh to its 
receptor opens 
the channel 


At the NMJ, the axon terminal releases 
ACh. The motor end-plate is the location 
of the ACh-receptors in the muscle fiber 

sarcolemma. When ACh molecules are 

released, they diffuse across a minute 
space called the synaptic cleft and bind to 
the receptors. 


The motor neurons that tell the skeletal muscle fibers to contract originate 
in the spinal cord, with a smaller number located in the brainstem for 
activation of skeletal muscles of the face, head, and neck. These neurons 
have long processes, called axons, which are specialized to transmit action 
potentials long distances— in this case, all the way from the spinal cord to 
the muscle itself (which may be up to three feet away). The axons of 
multiple neurons bundle together to form nerves, like wires bundled 
together in a cable. 


Signaling begins when a neuronal action potential travels along the axon 
of a motor neuron, and then along the individual branches to terminate at 
the NMJ. At the NMJ, the axon terminal releases a chemical messenger, or 
neurotransmitter, called acetylcholine (ACh). The ACh molecules diffuse 
across a minute space called the synaptic cleft and bind to ACh receptors 
located within the motor end-plate of the sarcolemma on the other side of 
the synapse. Once ACh binds, a channel in the ACh receptor opens and 
positively charged ions can pass through into the muscle fiber, causing it to 
depolarize, meaning that the membrane potential of the muscle fiber 
becomes less negative (closer to zero.) 


As the membrane depolarizes, another set of ion channels called voltage- 
gated sodium channels are triggered to open. Sodium ions enter the 
muscle fiber, and an action potential rapidly spreads (or “fires”) along the 
entire membrane to initiate excitation-contraction coupling. 


Things happen very quickly in the world of excitable membranes (just think 
about how quickly you can snap your fingers as soon as you decide to do 
it). Immediately following depolarization of the membrane, it repolarizes, 
re-establishing the negative membrane potential. Meanwhile, the ACh in 
the synaptic cleft is degraded by the enzyme acetylcholinesterase (AChE) 
so that the ACh cannot rebind to a receptor and reopen its channel, which 
would cause unwanted extended muscle excitation and contraction. 


Propagation of an action potential along the sarcolemma is the excitation 
portion of excitation-contraction coupling. Recall that this excitation 
actually triggers the release of calcium ions (Ca**) from its storage in the 


cell’s SR. For the action potential to reach the membrane of the SR, there 
are periodic invaginations in the sarcolemma, called T-tubules (“T” stands 
for “transverse”). You will recall that the diameter of a muscle fiber can be 
up to 100 pm, so these T-tubules ensure that the membrane can get close to 
the SR in the sarcoplasm. The arrangement of a T-tubule with the 
membranes of SR on either side is called a triad ([link]). The triad 
surrounds the cylindrical structure called a myofibril, which contains actin 
and myosin. 

The T-tubule 


Sarcolemma 


Sarcoplasmic 
reticulum 


Terminal cisternae 


Narrow T-tubules permit the 
conduction of electrical impulses. The 
SR functions to regulate intracellular 
levels of calcium. Two terminal 
cisternae (where enlarged SR connects 
to the T-tubule) and one T-tubule 
comprise a triad—a “threesome” of 
membranes, with those of SR on two 
sides and the T-tubule sandwiched 
between them. 


The T-tubules carry the action potential into the interior of the cell, which 
triggers the opening of calcium channels in the membrane of the adjacent 
SR, causing Ca** to diffuse out of the SR and into the sarcoplasm. It is the 


arrival of Ca** in the sarcoplasm that initiates contraction of the muscle 
fiber by its contractile units, or sarcomeres. 


Chapter Review 


Skeletal muscles contain connective tissue, blood vessels, and nerves. There 
are three layers of connective tissue: epimysium, perimysium, and 
endomysium. Skeletal muscle fibers are organized into groups called 
fascicles. Blood vessels and nerves enter the connective tissue and branch 
in the cell. Muscles attach to bones directly or through tendons or 
aponeuroses. Skeletal muscles maintain posture, stabilize bones and joints, 
control internal movement, and generate heat. 


Skeletal muscle fibers are long, multinucleated cells. The membrane of the 
cell is the sarcolemma; the cytoplasm of the cell is the sarcoplasm. The 
sarcoplasmic reticulum (SR) is a form of endoplasmic reticulum. Muscle 
fibers are composed of myofibrils. The striations are created by the 
organization of actin and myosin resulting in the banding pattern of 
myofibrils. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to learn more about macro- and microstructures of 
skeletal muscles. (a) What are the names of the “junction points” 
between sarcomeres? (b) What are the names of the “subunits” within 
the myofibrils that run the length of skeletal muscle fibers? (c) What is 
the “double strand of pearls” described in the video? (d) What gives a 
skeletal muscle fiber its striated appearance? 


Solution: 


(a) Z-lines. (b) Sarcomeres. (c) This is the arrangement of the actin and 
myosin filaments in a sarcomere. (d) The alternating strands of actin 


and myosin filaments. 
Exercise: 


Problem: 


Every skeletal muscle fiber is supplied by a motor neuron at the NMJ. 
Watch this video to learn more about what happens at the 
neuromuscular junction. (a) What is the definition of a motor unit? (b) 
What is the structural and functional difference between a large motor 
unit and a small motor unit? Can you give an example of each? (c) 
Why is the neurotransmitter acetylcholine degraded after binding to its 
receptor? 


Solution: 


(a) It is the number of skeletal muscle fibers supplied by a single motor 
neuron. (b) A large motor unit has one neuron supplying many skeletal 
muscle fibers for gross movements, like the Temporalis muscle, where 
1000 fibers are supplied by one neuron. A small motor has one neuron 
supplying few skeletal muscle fibers for very fine movements, like the 
extraocular eye muscles, where six fibers are supplied by one neuron. 
(c) To avoid prolongation of muscle contraction. 


Review Questions 


Exercise: 


Problem: 


The correct order for the smallest to the largest unit of organization in 
muscle tissue is 


a. fascicle, filament, muscle fiber, myofibril 
b. filament, myofibril, muscle fiber, fascicle 
c. muscle fiber, fascicle, filament, myofibril 
d. myofibril, muscle fiber, filament, fascicle 


Solution: 


B 


Exercise: 


Problem: Depolarization of the sarcolemma means 


a. the inside of the membrane has become less negative as sodium 
ions accumulate 

b. the outside of the membrane has become less negative as sodium 
ions accumulate 

c. the inside of the membrane has become more negative as sodium 
ions accumulate 

d. the sarcolemma has completely lost any electrical charge 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: 


What would happen to skeletal muscle if the epimysium were 
destroyed? 


Solution: 


Muscles would lose their integrity during powerful movements, 
resulting in muscle damage. 


Exercise: 


Problem: Describe how tendons facilitate body movement. 


Solution: 


When a muscle contracts, the force of movement is transmitted 
through the tendon, which pulls on the bone to produce skeletal 
movement. 


Exercise: 


Problem: What are the five primary functions of skeletal muscle? 


Solution: 


Produce movement of the skeleton, maintain posture and body 
position, support soft tissues, encircle openings of the digestive, 
urinary, and other tracts, and maintain body temperature. 


Exercise: 
Problem: 


What are the opposite roles of voltage-gated sodium channels and 
voltage-gated potassium channels? 


Solution: 


The opening of voltage-gated sodium channels, followed by the influx 
of Na’, transmits an Action Potential after the membrane has 
sufficiently depolarized. The delayed opening of potassium channels 
allows K" to exit the cell, to repolarize the membrane. 


Glossary 


acetylcholine (ACh) 
neurotransmitter that binds at a motor end-plate to trigger 
depolarization 


actin 


protein that makes up most of the thin myofilaments in a sarcomere 
muscle fiber 


action potential 
change in voltage of a cell membrane in response to a stimulus that 
results in transmission of an electrical signal; unique to neurons and 
muscle fibers 


aponeurosis 
broad, tendon-like sheet of connective tissue that attaches a skeletal 
muscle to another skeletal muscle or to a bone 


depolarize 
to reduce the voltage difference between the inside and outside of a 
cell’s plasma membrane (the sarcolemma for a muscle fiber), making 
the inside less negative than at rest 


endomysium 
loose, and well-hydrated connective tissue covering each muscle fiber 
in a skeletal muscle 


epimysium 
outer layer of connective tissue around a skeletal muscle 


excitation-contraction coupling 
sequence of events from motor neuron signaling to a skeletal muscle 
fiber to contraction of the fiber’s sarcomeres 


fascicle 
bundle of muscle fibers within a skeletal muscle 


motor end-plate 
sarcolemma of muscle fiber at the neuromuscular junction, with 
receptors for the neurotransmitter acetylcholine 


myofibril 
long, cylindrical organelle that runs parallel within the muscle fiber 
and contains the sarcomeres 


myosin 
protein that makes up most of the thick cylindrical myofilament within 
a sarcomere muscle fiber 


neuromuscular junction (NMJ) 
synapse between the axon terminal of a motor neuron and the section 
of the membrane of a muscle fiber with receptors for the acetylcholine 
released by the terminal 


neurotransmitter 
signaling chemical released by nerve terminals that bind to and 
activate receptors on target cells 


perimysium 
connective tissue that bundles skeletal muscle fibers into fascicles 
within a skeletal muscle 


sarcomere 
longitudinally, repeating functional unit of skeletal muscle, with all of 
the contractile and associated proteins involved in contraction 


sarcolemma 
plasma membrane of a skeletal muscle fiber 


sarcoplasm 
cytoplasm of a muscle cell 


sarcoplasmic reticulum (SR) 
specialized smooth endoplasmic reticulum, which stores, releases, and 
retrieves Ca** 


synaptic cleft 
space between a nerve (axon) terminal and a motor end-plate 


T-tubule 
projection of the sarcolemma into the interior of the cell 


thick filament 


the thick myosin strands and their multiple heads projecting from the 
center of the sarcomere toward, but not all to way to, the Z-discs 


thin filament 
thin strands of actin and its troponin-tropomyosin complex projecting 
from the Z-discs toward the center of the sarcomere 


triad 
the grouping of one T-tubule and two terminal cisternae 


troponin 
regulatory protein that binds to actin, tropomyosin, and calcium 


tropomyosin 
regulatory protein that covers myosin-binding sites to prevent actin 
from binding to myosin 


voltage-gated sodium channels 
membrane proteins that open sodium channels in response to a 
sufficient voltage change, and initiate and transmit the action potential 
as Na* enters through the channel 


Types of Muscle Fibers 
By the end of this section, you will be able to: 


e Describe the types of skeletal muscle fibers 
e Explain fast and slow muscle fibers 


Two criteria to consider when classifying the types of muscle fibers are how 
fast some fibers contract relative to others, and how fibers produce ATP. 
Using these criteria, there are three main types of skeletal muscle fibers. 
Slow oxidative (SO) fibers contract relatively slowly and use aerobic 
respiration (oxygen and glucose) to produce ATP. Fast oxidative (FO) 
fibers have fast contractions and primarily use aerobic respiration, but 
because they may switch to anaerobic respiration (glycolysis), can fatigue 
more quickly than SO fibers. Lastly, fast glycolytic (FG) fibers have fast 
contractions and primarily use anaerobic glycolysis. The FG fibers fatigue 
more quickly than the others. Most skeletal muscles in a human contain(s) 
all three types, although in varying proportions. 


The speed of contraction is dependent on how quickly myosin’s ATPase 
hydrolyzes ATP to produce cross-bridge action. Fast fibers hydrolyze ATP 
approximately twice as quickly as slow fibers, resulting in much quicker 
cross-bridge cycling (which pulls the thin filaments toward the center of the 
sarcomeres at a faster rate). The primary metabolic pathway used by a 
muscle fiber determines whether the fiber is classified as oxidative or 
glycolytic. If a fiber primarily produces ATP through aerobic pathways it is 
oxidative. More ATP can be produced during each metabolic cycle, making 
the fiber more resistant to fatigue. Glycolytic fibers primarily create ATP 
through anaerobic glycolysis, which produces less ATP per cycle. As a 
result, glycolytic fibers fatigue at a quicker rate. 


The oxidative fibers contain many more mitochondria than the glycolytic 
fibers, because aerobic metabolism, which uses oxygen (O>) in the 
metabolic pathway, occurs in the mitochondria. The SO fibers possess a 
large number of mitochondria and are capable of contracting for longer 
periods because of the large amount of ATP they can produce, but they have 
a relatively small diameter and do not produce a large amount of tension. 
SO fibers are extensively supplied with blood capillaries to supply Oz from 
the red blood cells in the bloodstream. The SO fibers also possess 


myoglobin, an O»-carrying molecule similar to Oj-carrying hemoglobin in 
the red blood cells. The myoglobin stores some of the needed O> within the 
fibers themselves (and gives SO fibers their red color). All of these features 
allow SO fibers to produce large quantities of ATP, which can sustain 
muscle activity without fatiguing for long periods of time. 


The fact that SO fibers can function for long periods without fatiguing 
makes them useful in maintaining posture, producing isometric 
contractions, stabilizing bones and joints, and making small movements that 
happen often but do not require large amounts of energy. They do not 
produce high tension, and thus they are not used for powerful, fast 
movements that require high amounts of energy and rapid cross-bridge 
cycling. 


FO fibers are sometimes called intermediate fibers because they possess 
characteristics that are intermediate between fast fibers and slow fibers. 
They produce ATP relatively quickly, more quickly than SO fibers, and thus 
can produce relatively high amounts of tension. They are oxidative because 
they produce ATP aerobically, possess high amounts of mitochondria, and 
do not fatigue quickly. However, FO fibers do not possess significant 
myoglobin, giving them a lighter color than the red SO fibers. FO fibers are 
used primarily for movements, such as walking, that require more energy 
than postural control but less energy than an explosive movement, such as 
sprinting. FO fibers are useful for this type of movement because they 
produce more tension than SO fibers but they are more fatigue-resistant 
than FG fibers. 


FG fibers primarily use anaerobic glycolysis as their ATP source. They 
have a large diameter and possess high amounts of glycogen, which is used 
in glycolysis to generate ATP quickly to produce high levels of tension. 
Because they do not primarily use aerobic metabolism, they do not possess 
substantial numbers of mitochondria or significant amounts of myoglobin 
and therefore have a white color. FG fibers are used to produce rapid, 
forceful contractions to make quick, powerful movements. These fibers 
fatigue quickly, permitting them to only be used for short periods. Most 
muscles possess a mixture of each fiber type. The predominant fiber type in 
a muscle is determined by the primary function of the muscle. 


Chapter Review 


ATP provides the energy for muscle contraction. The three mechanisms for 
ATP regeneration are creatine phosphate, anaerobic glycolysis, and aerobic 
metabolism. Creatine phosphate provides about the first 15 seconds of ATP 
at the beginning of muscle contraction. Anaerobic glycolysis produces 
small amounts of ATP in the absence of oxygen for a short period. Aerobic 
metabolism utilizes oxygen to produce much more ATP, allowing a muscle 
to work for longer periods. Muscle fatigue, which has many contributing 
factors, occurs when muscle can no longer contract. An oxygen debt is 
created as a result of muscle use. The three types of muscle fiber are slow 
oxidative (SO), fast oxidative (FO) and fast glycolytic (FG). SO fibers use 
aerobic metabolism to produce low power contractions over long periods 
and are slow to fatigue. FO fibers use aerobic metabolism to produce ATP 
but produce higher tension contractions than SO fibers. FG fibers use 
anaerobic metabolism to produce powerful, high-tension contractions but 
fatigue quickly. 


Review Questions 


Exercise: 


Problem: Muscle fatigue is caused by 


a. buildup of ATP and lactic acid levels 

b. exhaustion of energy reserves and buildup of lactic acid levels 

c. buildup of ATP and pyruvic acid levels 

d. exhaustion of energy reserves and buildup of pyruvic acid levels 


Solution: 


B 


Exercise: 


Problem: 


A sprinter would experience muscle fatigue sooner than a marathon 
runner due to 


a. anaerobic metabolism in the muscles of the sprinter 

b. anaerobic metabolism in the muscles of the marathon runner 
c. aerobic metabolism in the muscles of the sprinter 

d. glycolysis in the muscles of the marathon runner 


Solution: 


A 
Exercise: 


Problem: 


What aspect of creatine phosphate allows it to supply energy to 
muscles? 


a. ATPase activity 

b. phosphate bonds 
c. carbon bonds 

d. hydrogen bonds 


Solution: 


B 
Exercise: 


Problem: 


Drug X blocks ATP regeneration from ADP and phosphate. How will 
muscle cells respond to this drug? 


a. by absorbing ATP from the bloodstream 


b. by using ADP as an energy source 
c. by using glycogen as an energy source 
d. none of the above 


Solution: 


D 


Critical Thinking Questions 


Exercise: 
Problem: 


Why do muscle cells use creatine phosphate instead of glycolysis to 
supply ATP for the first few seconds of muscle contraction? 


Solution: 


Creatine phosphate is used because creatine phosphate and ADP are 
converted very quickly into ATP by creatine kinase. Glycolysis cannot 
generate ATP as quickly as creatine phosphate. 


Exercise: 


Problem: 


Is aerobic respiration more or less efficient than glycolysis? Explain 
your answer. 


Solution: 
Aerobic respiration is much more efficient than anaerobic glycolysis, 


yielding 36 ATP per molecule of glucose, as opposed to two ATP 
produced by glycolysis. 


Glossary 


fast glycolytic (FG) 
muscle fiber that primarily uses anaerobic glycolysis 


fast oxidative (FO) 
intermediate muscle fiber that is between slow oxidative and fast 
glycolytic fibers 


slow oxidative (SO) 
muscle fiber that primarily uses aerobic respiration 


Exercise and Muscle Performance 
By the end of this section, you will be able to: 


e Describe hypertrophy and atrophy 
e Explain how resistance exercise builds muscle 
e Explain how performance-enhancing substances affect muscle 


Physical training alters the appearance of skeletal muscles and can produce 
changes in muscle performance. Conversely, a lack of use can result in 
decreased performance and muscle appearance. Although muscle cells can 
change in size, new cells are not formed when muscles grow. Instead, 
structural proteins are added to muscle fibers in a process called 
hypertrophy, so cell diameter increases. The reverse, when structural 
proteins are lost and muscle mass decreases, is called atrophy. Age-related 
muscle atrophy is called sarcopenia. Cellular components of muscles can 
also undergo changes in response to changes in muscle use. 


Endurance Exercise 


Slow fibers are predominantly used in endurance exercises that require little 
force but involve numerous repetitions. The aerobic metabolism used by 
slow-twitch fibers allows them to maintain contractions over long periods. 
Endurance training modifies these slow fibers to make them even more 
efficient by producing more mitochondria to enable more aerobic 
metabolism and more ATP production. Endurance exercise can also 
increase the amount of myoglobin in a cell, as increased aerobic respiration 
increases the need for oxygen. Myoglobin is found in the sarcoplasm and 
acts as an oxygen storage supply for the mitochondria. 


The training can trigger the formation of more extensive capillary networks 
around the fiber, a process called angiogenesis, to supply oxygen and 
remove metabolic waste. To allow these capillary networks to supply the 
deep portions of the muscle, muscle mass does not greatly increase in order 
to maintain a smaller area for the diffusion of nutrients and gases. All of 
these cellular changes result in the ability to sustain low levels of muscle 
contractions for greater periods without fatiguing. 


The proportion of SO muscle fibers in muscle determines the suitability of 
that muscle for endurance, and may benefit those participating in endurance 
activities. Postural muscles have a large number of SO fibers and relatively 
few FO and FG fibers, to keep the back straight ({link]). Endurance athletes, 
like marathon-runners also would benefit from a larger proportion of SO 
fibers, but it is unclear if the most-successful marathoners are those with 
naturally high numbers of SO fibers, or whether the most successful 
marathon runners develop high numbers of SO fibers with repetitive 
training. Endurance training can result in overuse injuries such as stress 
fractures and joint and tendon inflammation. 
Marathoners 

a 


Long-distance runners have a large 
number of SO fibers and relatively few 
FO and FG fibers. (credit: 
“Tseo2”/Wikimedia Commons) 


Resistance Exercise 


Resistance exercises, as opposed to endurance exercise, require large 
amounts of FG fibers to produce short, powerful movements that are not 
repeated over long periods. The high rates of ATP hydrolysis and cross- 
bridge formation in FG fibers result in powerful muscle contractions. 


Muscles used for power have a higher ratio of FG to SO/FO fibers, and 
trained athletes possess even higher levels of FG fibers in their muscles. 
Resistance exercise affects muscles by increasing the formation of 
myofibrils, thereby increasing the thickness of muscle fibers. This added 
structure causes hypertrophy, or the enlargement of muscles, exemplified by 
the large skeletal muscles seen in body builders and other athletes ([link]). 
Because this muscular enlargement is achieved by the addition of structural 
proteins, athletes trying to build muscle mass often ingest large amounts of 
protein. 

Hypertrophy 


Body builders have a large number of 
FG fibers and relatively few FO and 
SO fibers. (credit: Lin Mei/flickr) 


Except for the hypertrophy that follows an increase in the number of 
sarcomeres and myofibrils in a skeletal muscle, the cellular changes 
observed during endurance training do not usually occur with resistance 
training. There is usually no significant increase in mitochondria or 
capillary density. However, resistance training does increase the 
development of connective tissue, which adds to the overall mass of the 
muscle and helps to contain muscles as they produce increasingly powerful 
contractions. Tendons also become stronger to prevent tendon damage, as 
the force produced by muscles is transferred to tendons that attach the 
muscle to bone. 


For effective strength training, the intensity of the exercise must continually 
be increased. For instance, continued weight lifting without increasing the 
weight of the load does not increase muscle size. To produce ever-greater 
results, the weights lifted must become increasingly heavier, making it more 
difficult for muscles to move the load. The muscle then adapts to this 
heavier load, and an even heavier load must be used if even greater muscle 
mass is desired. 


If done improperly, resistance training can lead to overuse injuries of the 
muscle, tendon, or bone. These injuries can occur if the load is too heavy or 
if the muscles are not given sufficient time between workouts to recover or 
if joints are not aligned properly during the exercises. Cellular damage to 
muscle fibers that occurs after intense exercise includes damage to the 
sarcolemma and myofibrils. This muscle damage contributes to the feeling 
of soreness after strenuous exercise, but muscles gain mass as this damage 
is repaired, and additional structural proteins are added to replace the 
damaged ones. Overworking skeletal muscles can also lead to tendon 
damage and even skeletal damage if the load is too great for the muscles to 
bear. 


Performance-Enhancing Substances 


Some athletes attempt to boost their performance by using various agents 
that may enhance muscle performance. Anabolic steroids are one of the 
more widely known agents used to boost muscle mass and increase power 
output. Anabolic steroids are a form of testosterone, a male sex hormone 
that stimulates muscle formation, leading to increased muscle mass. 


Endurance athletes may also try to boost the availability of oxygen to 
muscles to increase aerobic respiration by using substances such as 
erythropoietin (EPO), a hormone normally produced in the kidneys, which 
triggers the production of red blood cells. The extra oxygen carried by these 
blood cells can then be used by muscles for aerobic respiration. Human 
growth hormone (hGH) is another supplement, and although it can facilitate 
building muscle mass, its main role is to promote the healing of muscle and 
other tissues after strenuous exercise. Increased hGH may allow for faster 


recovery after muscle damage, reducing the rest required after exercise, and 
allowing for more sustained high-level performance. 


Although performance-enhancing substances often do improve 
performance, most are banned by governing bodies in sports and are illegal 
for nonmedical purposes. Their use to enhance performance raises ethical 
issues of cheating because they give users an unfair advantage over 
nonusers. A greater concern, however, is that their use carries serious health 
risks. The side effects of these substances are often significant, 
nonreversible, and in some cases fatal. The physiological strain caused by 
these substances is often greater than what the body can handle, leading to 
effects that are unpredictable and dangerous. Anabolic steroid use has been 
linked to infertility, aggressive behavior, cardiovascular disease, and brain 
cancer. 


Similarly, some athletes have used creatine to increase power output. 
Creatine phosphate provides quick bursts of ATP to muscles in the initial 
stages of contraction. Increasing the amount of creatine available to cells is 
thought to produce more ATP and therefore increase explosive power 
output, although its effectiveness as a supplement has been questioned. 


Note: 

Everyday Connection 

Aging and Muscle Tissue 

Although atrophy due to disuse can often be reversed with exercise, muscle 
atrophy with age, referred to as sarcopenia, is irreversible. This is a 
primary reason why even highly trained athletes succumb to declining 
performance with age. This decline is noticeable in athletes whose sports 
require strength and powerful movements, such as sprinting, whereas the 
effects of age are less noticeable in endurance athletes such as marathon 
runners or long-distance cyclists. As muscles age, muscle fibers die, and 
they are replaced by connective tissue and adipose tissue ([link]). Because 
those tissues cannot contract and generate force as muscle can, muscles 
lose the ability to produce powerful contractions. The decline in muscle 
mass causes a loss of strength, including the strength required for posture 
and mobility. This may be caused by a reduction in FG fibers that 


hydrolyze ATP quickly to produce short, powerful contractions. Muscles in 
older people sometimes possess greater numbers of SO fibers, which are 
responsible for longer contractions and do not produce powerful 
movements. There may also be a reduction in the size of motor units, 
resulting in fewer fibers being stimulated and less muscle tension being 
produced. 
Atrophy 

{ ) 


Atrophied 
Muscle 


Normal 
Muscle 


Muscle mass is reduced as muscles 
atrophy with disuse. 


Sarcopenia can be delayed to some extent by exercise, as training adds 
structural proteins and causes cellular changes that can offset the effects of 
atrophy. Increased exercise can produce greater numbers of cellular 
mitochondria, increase capillary density, and increase the mass and 
strength of connective tissue. The effects of age-related atrophy are 
especially pronounced in people who are sedentary, as the loss of muscle 
cells is displayed as functional impairments such as trouble with 
locomotion, balance, and posture. This can lead to a decrease in quality of 
life and medical problems, such as joint problems because the muscles that 
stabilize bones and joints are weakened. Problems with locomotion and 
balance can also cause various injuries due to falls. 


Chapter Review 


Hypertrophy is an increase in muscle mass due to the addition of structural 
proteins. The opposite of hypertrophy is atrophy, the loss of muscle mass 
due to the breakdown of structural proteins. Endurance exercise causes an 
increase in cellular mitochondria, myoglobin, and capillary networks in SO 
fibers. Endurance athletes have a high level of SO fibers relative to the 
other fiber types. Resistance exercise causes hypertrophy. Power-producing 
muscles have a higher number of FG fibers than of slow fibers. Strenuous 
exercise causes muscle cell damage that requires time to heal. Some 
athletes use performance-enhancing substances to enhance muscle 
performance. Muscle atrophy due to age is called sarcopenia and occurs as 
muscle fibers die and are replaced by connective and adipose tissue. 


Review Questions 


Exercise: 


Problem: 


The muscles of a professional sprinter are most likely to have 


a. 80 percent fast-twitch muscle fibers and 20 percent slow-twitch 
muscle fibers 

b. 20 percent fast-twitch muscle fibers and 80 percent slow-twitch 
muscle fibers 

c. 50 percent fast-twitch muscle fibers and 50 percent slow-twitch 
muscle fibers 

d. 40 percent fast-twitch muscle fibers and 60 percent slow-twitch 
muscle fibers 


Solution: 


A 


Exercise: 


Problem: 


The muscles of a professional marathon runner are most likely to have 


a. 80 percent fast-twitch muscle fibers and 20 percent slow-twitch 
muscle fibers 

b. 20 percent fast-twitch muscle fibers and 80 percent slow-twitch 
muscle fibers 

c. 50 percent fast-twitch muscle fibers and 50 percent slow-twitch 
muscle fibers 

d. 40 percent fast-twitch muscle fibers and 60 percent slow-twitch 
muscle fibers 


Solution: 
B 
Exercise: 
Problem: Which of the following statements is true? 


a. Fast fibers have a small diameter. 

b. Fast fibers contain loosely packed myofibrils. 
c. Fast fibers have large glycogen reserves. 

d. Fast fibers have many mitochondria. 


Solution: 
Cc 
Exercise: 
Problem: Which of the following statements is false? 


a. Slow fibers have a small network of capillaries. 


b. Slow fibers contain the pigment myoglobin. 
c. Slow fibers contain a large number of mitochondria. 
d. Slow fibers contract for extended periods. 


Solution: 


A 


Critical Thinking Questions 


Exercise: 
Problem: 


What changes occur at the cellular level in response to endurance 
training? 


Solution: 


Endurance training modifies slow fibers to make them more efficient 
by producing more mitochondria to enable more aerobic metabolism 
and more ATP production. Endurance exercise can also increase the 
amount of myoglobin in a cell and formation of more extensive 
capillary networks around the fiber. 


Exercise: 


Problem: 


What changes occur at the cellular level in response to resistance 
training? 


Solution: 


Resistance exercises affect muscles by causing the formation of more 
actin and myosin, increasing the structure of muscle fibers. 


Glossary 


angiogenesis 
formation of blood capillary networks 


atrophy 
loss of structural proteins from muscle fibers 


hypertrophy 
addition of structural proteins to muscle fibers 


Sarcopenia 
age-related muscle atrophy 


Cardiac Muscle Tissue 
By the end of this section, you will be able to: 


¢ Describe intercalated discs and gap junctions 
e Describe a desmosome 


Cardiac muscle tissue is only found in the heart. Highly coordinated 
contractions of cardiac muscle pump blood into the vessels of the 
circulatory system. Similar to skeletal muscle, cardiac muscle is striated and 
organized into sarcomeres, possessing the same banding organization as 
skeletal muscle ({link]). However, cardiac muscle fibers are shorter than 
skeletal muscle fibers and usually contain only one nucleus, which is 
located in the central region of the cell. Cardiac muscle fibers also possess 
many mitochondria and myoglobin, as ATP is produced primarily through 
aerobic metabolism. Cardiac muscle fibers cells also are extensively 
branched and are connected to one another at their ends by intercalated 
discs. An intercalated disc allows the cardiac muscle cells to contract in a 
wave-like pattern so that the heart can work as a pump. 

Cardiac Muscle Tissue 


Cardiac muscle tissue is only found in the 

heart. LM x 1600. (Micrograph provided 

by the Regents of University of Michigan 
Medical School © 2012) 


Note: 


== 
mmm ~OPENStaX COLLEGE’ 
— 
— 

. a 
. 

Dy) 

r | 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. 


Intercalated discs are part of the sarcolemma and contain two structures 
important in cardiac muscle contraction: gap junctions and desmosomes. A 
gap junction forms channels between adjacent cardiac muscle fibers that 
allow the depolarizing current produced by cations to flow from one cardiac 
muscle cell to the next. This joining is called electric coupling, and in 
cardiac muscle it allows the quick transmission of action potentials and the 
coordinated contraction of the entire heart. This network of electrically 
connected cardiac muscle cells creates a functional unit of contraction 
called a syncytium. The remainder of the intercalated disc is composed of 
desmosomes. A desmosome is a cell structure that anchors the ends of 
cardiac muscle fibers together so the cells do not pull apart during the stress 
of individual fibers contracting ([link]). 

Cardiac Muscle 


Capillary Intercalated discs 


Desmosome 


Gap junction 


Nucleus Cardiac 
muscle fiber 


Intercalated discs are part of the cardiac muscle 
sarcolemma and they contain gap junctions and 
desmosomes. 


Contractions of the heart (heartbeats) are controlled by specialized cardiac 
muscle cells called pacemaker cells that directly control heart rate. 
Although cardiac muscle cannot be consciously controlled, the pacemaker 
cells respond to signals from the autonomic nervous system (ANS) to speed 
up or slow down the heart rate. The pacemaker cells can also respond to 
various hormones that modulate heart rate to control blood pressure. 


The wave of contraction that allows the heart to work as a unit, called a 
functional syncytium, begins with the pacemaker cells. This group of cells 
is self-excitable and able to depolarize to threshold and fire action potentials 
on their own, a feature called autorhythmicity; they do this at set intervals 
which determine heart rate. Because they are connected with gap junctions 
to surrounding muscle fibers and the specialized fibers of the heart’s 
conduction system, the pacemaker cells are able to transfer the 
depolarization to the other cardiac muscle fibers in a manner that allows the 
heart to contract in a coordinated manner. 


Another feature of cardiac muscle is its relatively long action potentials in 
its fibers, having a sustained depolarization “plateau.” The plateau is 
produced by Ca** entry though voltage-gated calcium channels in the 
sarcolemma of cardiac muscle fibers. This sustained depolarization (and 
Ca** entry) provides for a longer contraction than is produced by an action 
potential in skeletal muscle. Unlike skeletal muscle, a large percentage of 
the Ca™* that initiates contraction in cardiac muscles comes from outside 
the cell rather than from the SR. 


Chapter Review 


Cardiac muscle is striated muscle that is present only in the heart. Cardiac 
muscle fibers have a single nucleus, are branched, and joined to one another 
by intercalated discs that contain gap junctions for depolarization between 


cells and desmosomes to hold the fibers together when the heart contracts. 
Contraction in each cardiac muscle fiber is triggered by Ca™ ions in a 
similar manner as skeletal muscle, but here the Ca** ions come from SR 
and through voltage-gated calcium channels in the sarcolemma. Pacemaker 
cells stimulate the spontaneous contraction of cardiac muscle as a 
functional unit, called a syncytium. 


Review Questions 


Exercise: 


Problem: 
Cardiac muscles differ from skeletal muscles in that they 


a. are striated 

b. utilize aerobic metabolism 
c. contain myofibrils 

d. contain intercalated discs 


Solution: 


D 
Exercise: 
Problem: 


If cardiac muscle cells were prevented from undergoing aerobic 
metabolism, they ultimately would 


a. undergo glycolysis 
b. synthesize ATP 
c. stop contracting 
d. start contracting 


Solution: 


Critical Thinking Questions 


Exercise: 
Problem: 


What would be the drawback of cardiac contractions being the same 
duration as skeletal muscle contractions? 


Solution: 


An action potential could reach a cardiac muscle cell before it has 
entered the relaxation phase, resulting in the sustained contractions of 
tetanus. If this happened, the heart would not beat regularly. 


Exercise: 
Problem: 


How are cardiac muscle cells similar to and different from skeletal 
muscle cells? 


Solution: 


Cardiac and skeletal muscle cells both contain ordered myofibrils and 
are striated. Cardiac muscle cells are branched and contain intercalated 
discs, which skeletal muscles do not have. 


Glossary 


autorhythmicity 
heart’s ability to control its own contractions 


desmosome 
cell structure that anchors the ends of cardiac muscle fibers to allow 
contraction to occur 


intercalated disc 
part of the sarcolemma that connects cardiac tissue, and contains gap 
junctions and desmosomes 


Smooth Muscle 
By the end of this section, you will be able to: 


e Describe a dense body 

e Explain how smooth muscle works with internal organs and 
passageways through the body 

e Explain how smooth muscles differ from skeletal and cardiac muscles 

e Explain the difference between single-unit and multi-unit smooth 
muscle 


Smooth muscle (so-named because the cells do not have striations) is 
present in the walls of hollow organs like the urinary bladder, uterus, 
stomach, intestines, and in the walls of passageways, such as the arteries 
and veins of the circulatory system, and the tracts of the respiratory, urinary, 
and reproductive systems ({link]ab). Smooth muscle is also present in the 
eyes, where it functions to change the size of the iris and alter the shape of 
the lens; and in the skin where it causes hair to stand erect in response to 
cold temperature or fear. 

Smooth Muscle Tissue 


Autonomic 
neurons 


Nucleus 


Muscle 
fibers 


(a) 


(b) 


Smooth muscle tissue is found around organs in the digestive, 
respiratory, reproductive tracts and the iris of the eye. LM x 
1600. (Micrograph provided by the Regents of University of 

Michigan Medical School © 2012) 


Note: 


— 7 
mess Openstax COLLEGE 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. 


Smooth muscle fibers are spindle-shaped (wide in the middle and tapered at 
both ends, somewhat like a football) and have a single nucleus; they range 
from about 30 to 200 pm (thousands of times shorter than skeletal muscle 
fibers), and they produce their own connective tissue, endomysium. 
Although they do not have striations and sarcomeres, smooth muscle fibers 
do have actin and myosin contractile proteins, and thick and thin filaments. 
These thin filaments are anchored by dense bodies. A dense body is 
analogous to the Z-discs of skeletal and cardiac muscle fibers and is 
fastened to the sarcolemma. Calcium ions are supplied by the SR in the 
fibers and by sequestration from the extracellular fluid through membrane 
indentations called calveoli. 


Because smooth muscle cells do not contain troponin, cross-bridge 
formation is not regulated by the troponin-tropomyosin complex but instead 
by the regulatory protein calmodulin. In a smooth muscle fiber, external 
Ca™* ions passing through opened calcium channels in the sarcolemma, and 
additional Ca** released from SR, bind to calmodulin. The Ca**- 
calmodulin complex then activates an enzyme called myosin (light chain) 
kinase, which, in turn, activates the myosin heads by phosphorylating them 
(converting ATP to ADP and P;, with the P; attaching to the head). The 
heads can then attach to actin-binding sites and pull on the thin filaments. 
The thin filaments also are anchored to the dense bodies; the structures 
invested in the inner membrane of the sarcolemma (at adherens junctions) 
that also have cord-like intermediate filaments attached to them. When the 
thin filaments slide past the thick filaments, they pull on the dense bodies, 
structures tethered to the sarcolemma, which then pull on the intermediate 
filaments networks throughout the sarcoplasm. This arrangement causes the 
entire muscle fiber to contract in a manner whereby the ends are pulled 
toward the center, causing the midsection to bulge in a corkscrew motion 
([link]). 

Muscle Contraction 


Relaxed muscle cell Contracted muscle cell 


—> = 


Intermediate filaments Dense bodies 


The dense bodies and intermediate filaments are networked 
through the sarcoplasm, which cause the muscle fiber to 
contract. 


Although smooth muscle contraction relies on the presence of Ca** ions, 
smooth muscle fibers have a much smaller diameter than skeletal muscle 
cells. T-tubules are not required to reach the interior of the cell and 
therefore not necessary to transmit an action potential deep into the fiber. 
Smooth muscle fibers have a limited calcium-storing SR but have calcium 
channels in the sarcolemma (similar to cardiac muscle fibers) that open 
during the action potential along the sarcolemma. The influx of 
extracellular Ca** ions, which diffuse into the sarcoplasm to reach the 
calmodulin, accounts for most of the Ca** that triggers contraction of a 
smooth muscle cell. 


Muscle contraction continues until ATP-dependent calcium pumps actively 
transport Ca** ions back into the SR and out of the cell. However, a low 
concentration of calcium remains in the sarcoplasm to maintain muscle 
tone. This remaining calcium keeps the muscle slightly contracted, which is 
important in certain tracts and around blood vessels. 


Because most smooth muscles must function for long periods without rest, 
their power output is relatively low, but contractions can continue without 
using large amounts of energy. Some smooth muscle can also maintain 
contractions even as Ca** is removed and myosin kinase is 
inactivated/dephosphorylated. This can happen as a subset of cross-bridges 
between myosin heads and actin, called latch-bridges, keep the thick and 
thin filaments linked together for a prolonged period, and without the need 
for ATP. This allows for the maintaining of muscle “tone” in smooth muscle 


that lines arterioles and other visceral organs with very little energy 
expenditure. 


Smooth muscle is not under voluntary control; thus, it is called involuntary 
muscle. The triggers for smooth muscle contraction include hormones, 
neural stimulation by the ANS, and local factors. In certain locations, such 
as the walls of visceral organs, stretching the muscle can trigger its 
contraction (the stress-relaxation response). 


Axons of neurons in the ANS do not form the highly organized NMJs with 
smooth muscle, as seen between motor neurons and skeletal muscle fibers. 
Instead, there is a series of neurotransmitter-filled bulges called varicosities 
as an axon courses through smooth muscle, loosely forming motor units 
({link]). A varicosity releases neurotransmitters into the synaptic cleft. 
Also, visceral muscle in the walls of the hollow organs (except the heart) 
contains pacesetter cells. A pacesetter cell can spontaneously trigger action 
potentials and contractions in the muscle. 

Motor Units 


Varicosity 


Vesicles with 
neurotransmitters . 
Autonomic neuron 


—— Smooth muscle cells 


A series of axon-like swelling, called varicosities or “boutons,” 
from autonomic neurons form motor units through the smooth 
muscle. 


Smooth muscle is organized in two ways: as single-unit smooth muscle, 
which is much more common; and as multiunit smooth muscle. The two 


types have different locations in the body and have different characteristics. 
Single-unit muscle has its muscle fibers joined by gap junctions so that the 
muscle contracts as a single unit. This type of smooth muscle is found in the 
walls of all visceral organs except the heart (which has cardiac muscle in its 
walls), and so it is commonly called visceral muscle. Because the muscle 
fibers are not constrained by the organization and stretchability limits of 
sarcomeres, visceral smooth muscle has a stress-relaxation response. This 
means that as the muscle of a hollow organ is stretched when it fills, the 
mechanical stress of the stretching will trigger contraction, but this is 
immediately followed by relaxation so that the organ does not empty its 
contents prematurely. This is important for hollow organs, such as the 
stomach or urinary bladder, which continuously expand as they fill. The 
smooth muscle around these organs also can maintain a muscle tone when 
the organ empties and shrinks, a feature that prevents “flabbiness” in the 
empty organ. In general, visceral smooth muscle produces slow, steady 
contractions that allow substances, such as food in the digestive tract, to 
move through the body. 


Multiunit smooth muscle cells rarely possess gap junctions, and thus are not 
electrically coupled. As a result, contraction does not spread from one cell 
to the next, but is instead confined to the cell that was originally stimulated. 
Stimuli for multiunit smooth muscles come from autonomic nerves or 
hormones but not from stretching. This type of tissue is found around large 
blood vessels, in the respiratory airways, and in the eyes. 


Hyperplasia in Smooth Muscle 


Similar to skeletal and cardiac muscle cells, smooth muscle can undergo 
hypertrophy to increase in size. Unlike other muscle, smooth muscle can 
also divide to produce more cells, a process called hyperplasia. This can 
most evidently be observed in the uterus at puberty, which responds to 
increased estrogen levels by producing more uterine smooth muscle fibers, 
and greatly increases the size of the myometrium. 


Sections Summary 


Smooth muscle is found throughout the body around various organs and 
tracts. Smooth muscle cells have a single nucleus, and are spindle-shaped. 
Smooth muscle cells can undergo hyperplasia, mitotically dividing to 
produce new cells. The smooth cells are nonstriated, but their sarcoplasm is 
filled with actin and myosin, along with dense bodies in the sarcolemma to 
anchor the thin filaments and a network of intermediate filaments involved 
in pulling the sarcolemma toward the fiber’s middle, shortening it in the 
process. Ca** ions trigger contraction when they are released from SR and 
enter through opened voltage-gated calcium channels. Smooth muscle 
contraction is initiated when the Ca‘™ binds to intracellular calmodulin, 
which then activates an enzyme called myosin kinase that phosphorylates 
myosin heads so they can form the cross-bridges with actin and then pull on 
the thin filaments. Smooth muscle can be stimulated by pacesetter cells, by 
the autonomic nervous system, by hormones, spontaneously, or by 
stretching. The fibers in some smooth muscle have latch-bridges, cross- 
bridges that cycle slowly without the need for ATP; these muscles can 
maintain low-level contractions for long periods. Single-unit smooth muscle 
tissue contains gap junctions to synchronize membrane depolarization and 
contractions so that the muscle contracts as a single unit. Single-unit 
smooth muscle in the walls of the viscera, called visceral muscle, has a 
stress-relaxation response that permits muscle to stretch, contract, and relax 
as the organ expands. Multiunit smooth muscle cells do not possess gap 
junctions, and contraction does not spread from one cell to the next. 


Multiple Choice 


Exercise: 


Problem: 


Smooth muscles differ from skeletal and cardiac muscles in that they 


a. lack myofibrils 

b. are under voluntary control 
c. lack myosin 

d. lack actin 


Solution: 


A 
Exercise: 


Problem: 
Which of the following statements describes smooth muscle cells? 


a. They are resistant to fatigue. 

b. They have a rapid onset of contractions. 

c. They cannot exhibit tetanus. 

d. They primarily use anaerobic metabolism. 


Solution: 


A 


Critical Thinking Questions 


Exercise: 
Problem: 


Why can smooth muscles contract over a wider range of resting 
lengths than skeletal and cardiac muscle? 


Solution: 


Smooth muscles can contract over a wider range of resting lengths 
because the actin and myosin filaments in smooth muscle are not as 
rigidly organized as those in skeletal and cardiac muscle. 


Exercise: 


Problem: 


Describe the differences between single-unit smooth muscle and 
multiunit smooth muscle. 


Solution: 


Single-unit smooth muscle is found in the walls of hollow organs; 
multiunit smooth muscle is found in airways to the lungs and large 
arteries. Single-unit smooth muscle cells contract synchronously, they 
are coupled by gap junctions, and they exhibit spontaneous action 
potential. Multiunit smooth cells lack gap junctions, and their 
contractions are not synchronous. 


Glossary 


calmodulin 
regulatory protein that facilitates contraction in smooth muscles 


dense body 
sarcoplasmic structure that attaches to the sarcolemma and shortens the 
muscle as thin filaments slide past thick filaments 


hyperplasia 
process in which one cell splits to produce new cells 


latch-bridges 
subset of a cross-bridge in which actin and myosin remain locked 
together 


pacesetter cell 
cell that triggers action potentials in smooth muscle 


stress-relaxation response 
relaxation of smooth muscle tissue after being stretched 


varicosity 


enlargement of neurons that release neurotransmitters into synaptic 
clefts 


visceral muscle 
smooth muscle found in the walls of visceral organs 


Development and Regeneration of Muscle Tissue 
By the end of this section, you will be able to: 


e Describe the function of satellite cells 
e Define fibrosis 
e Explain which muscle has the greatest regeneration ability 


Most muscle tissue of the body arises from embryonic mesoderm. Paraxial 
mesodermal cells adjacent to the neural tube form blocks of cells called 
somites. Skeletal muscles, excluding those of the head and limbs, develop 
from mesodermal somites, whereas skeletal muscle in the head and limbs 
develop from general mesoderm. Somites give rise to myoblasts. A 
myoblast is a muscle-forming stem cell that migrates to different regions in 
the body and then fuse(s) to form a syncytium, or myotube. As a myotube 
is formed from many different myoblast cells, it contains many nuclei, but 
has a continuous cytoplasm. This is why skeletal muscle cells are 
multinucleate, as the nucleus of each contributing myoblast remains intact 
in the mature skeletal muscle cell. However, cardiac and smooth muscle 
cells are not multinucleate because the myoblasts that form their cells do 
not fuse. 


Gap junctions develop in the cardiac and single-unit smooth muscle in the 
early stages of development. In skeletal muscles, ACh receptors are initially 
present along most of the surface of the myoblasts, but spinal nerve 
innervation causes the release of growth factors that stimulate the formation 
of motor end-plates and NMJs. As neurons become active, electrical signals 
that are sent through the muscle influence the distribution of slow and fast 
fibers in the muscle. 


Although the number of muscle cells is set during development, satellite 
cells help to repair skeletal muscle cells. A satellite cell is similar to a 
myoblast because it is a type of stem cell; however, satellite cells are 
incorporated into muscle cells and facilitate the protein synthesis required 
for repair and growth. These cells are located outside the sarcolemma and 
are stimulated to grow and fuse with muscle cells by growth factors that are 
released by muscle fibers under certain forms of stress. Satellite cells can 
regenerate muscle fibers to a very limited extent, but they primarily help to 
repair damage in living cells. If a cell is damaged to a greater extent than 


can be repaired by satellite cells, the muscle fibers are replaced by scar 
tissue in a process called fibrosis. Because scar tissue cannot contract, 
muscle that has sustained significant damage loses strength and cannot 
produce the same amount of power or endurance as it could before being 
damaged. 


Smooth muscle tissue can regenerate from a type of stem cell called a 
pericyte, which is found in some small blood vessels. Pericytes allow 
smooth muscle cells to regenerate and repair much more readily than 
skeletal and cardiac muscle tissue. Similar to skeletal muscle tissue, cardiac 
muscle does not regenerate to a great extent. Dead cardiac muscle tissue is 
replaced by scar tissue, which cannot contract. As scar tissue accumulates, 
the heart loses its ability to pump because of the loss of contractile power. 
However, some minor regeneration may occur due to stem cells found in 
the blood that occasionally enter cardiac tissue. 


Note: 

Career Connections 

Physical Therapist 

As muscle cells die, they are not regenerated but instead are replaced by 
connective tissue and adipose tissue, which do not possess the contractile 
abilities of muscle tissue. Muscles atrophy when they are not used, and 
over time if atrophy is prolonged, muscle cells die. It is therefore important 
that those who are susceptible to muscle atrophy exercise to maintain 
muscle function and prevent the complete loss of muscle tissue. In extreme 
cases, when movement is not possible, electrical stimulation can be 
introduced to a muscle from an external source. This acts as a substitute for 
endogenous neural stimulation, stimulating the muscle to contract and 
preventing the loss of proteins that occurs with a lack of use. 
Physiotherapists work with patients to maintain muscles. They are trained 
to target muscles susceptible to atrophy, and to prescribe and monitor 
exercises designed to stimulate those muscles. There are various causes of 
atrophy, including mechanical injury, disease, and age. After breaking a 
limb or undergoing surgery, muscle use is impaired and can lead to disuse 
atrophy. If the muscles are not exercised, this atrophy can lead to long-term 


muscle weakness. A stroke can also cause muscle impairment by 
interrupting neural stimulation to certain muscles. Without neural inputs, 
these muscles do not contract and thus begin to lose structural proteins. 
Exercising these muscles can help to restore muscle function and minimize 
functional impairments. Age-related muscle loss is also a target of physical 
therapy, as exercise can reduce the effects of age-related atrophy and 
improve muscle function. 

The goal of a physiotherapist is to improve physical functioning and 
reduce functional impairments; this is achieved by understanding the cause 
of muscle impairment and assessing the capabilities of a patient, after 
which a program to enhance these capabilities is designed. Some factors 
that are assessed include strength, balance, and endurance, which are 
continually monitored as exercises are introduced to track improvements in 
muscle function. Physiotherapists can also instruct patients on the proper 
use of equipment, such as crutches, and assess whether someone has 
sufficient strength to use the equipment and when they can function 
without it. 


Chapter Review 


Muscle tissue arises from embryonic mesoderm. Somites give rise to 
myoblasts and fuse to form a myotube. The nucleus of each contributing 
myoblast remains intact in the mature skeletal muscle cell, resulting in a 
mature, multinucleate cell. Satellite cells help to repair skeletal muscle 
cells. Smooth muscle tissue can regenerate from stem cells called pericytes, 
whereas dead cardiac muscle tissue is replaced by scar tissue. Aging causes 
muscle mass to decrease and be replaced by noncontractile connective 
tissue and adipose tissue. 


Review Questions 


Exercise: 


Problem: 


From which embryonic cell type does muscle tissue develop? 


a. ganglion cells 
b. myotube cells 
c. myoblast cells 
d. satellite cells 


Solution: 
C 
Exercise: 
Problem: Which cell type helps to repair injured muscle fibers? 


a. ganglion cells 
b. myotube cells 
c. myoblast cells 
d. satellite cells 


Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: 


Why is muscle that has sustained significant damage unable to produce 
the same amount of power as it could before being damaged? 


Solution: 


If the damage exceeds what can be repaired by satellite cells, the 
damaged tissue is replaced by scar tissue, which cannot contract. 


Exercise: 


Problem: 


Which muscle type(s) (skeletal, smooth, or cardiac) can regenerate 
new muscle cells/fibers? Explain your answer. 


Solution: 


Smooth muscle tissue can regenerate from stem cells called pericytes, 
cells found in some small blood vessels. These allow smooth muscle 
cells to regenerate and repair much more readily than skeletal and 
cardiac muscle tissue. 


Glossary 


fibrosis 
replacement of muscle fibers by scar tissue 


myoblast 
muscle-forming stem cell 


myotube 
fusion of many myoblast cells 


pericyte 
stem cell that regenerates smooth muscle cells 


satellite cell 
stem cell that helps to repair muscle cells 


somites 
blocks of paraxial mesoderm cells 


Interactions of Skeletal Muscles, Their Fascicle Arrangement, and Their 
Lever Systems 
By the end of this section, you will be able to: 


e Compare and contrast agonist and antagonist muscles 

e Describe how fascicles are arranged within a skeletal muscle 

e Explain the major events of a skeletal muscle contraction within a 
muscle in generating force 


To move the skeleton, the tension created by the contraction of the fibers in 
most skeletal muscles is transferred to the tendons. The tendons are strong 
bands of dense, regular connective tissue that connect muscles to bones. 
The bone connection is why this muscle tissue is called skeletal muscle. 


Interactions of Skeletal Muscles in the Body 


To pull on a bone, that is, to change the angle at its synovial joint, which 
essentially moves the skeleton, a skeletal muscle must also be attached to a 
fixed part of the skeleton. The moveable end of the muscle that attaches to 
the bone being pulled is called the muscle’s insertion, and the end of the 
muscle attached to a fixed (stabilized) bone is called the origin. During 
forearm flexion—bending the elbow—the brachioradialis assists the 
brachialis. 


Although a number of muscles may be involved in an action, the principal 
muscle involved is called the prime mover, or agonist. To lift a cup, a 
muscle called the biceps brachii is actually the prime mover; however, 
because it can be assisted by the brachialis, the brachialis is called a 
synergist in this action ([link]). A synergist can also be a fixator that 
stabilizes the bone that is the attachment for the prime mover’s origin. 
Prime Movers and Synergists 


Biceps brachii 
(prime mover) 


Brachioradialis 
(synergist) 


Biceps brachii 
(dissected) 


Brachialis 
(synergist) 


Brachioradialis 


The biceps brachii flex the lower arm. The 
brachoradialis, in the forearm, and brachialis, 
located deep to the biceps in the upper arm, are 
both synergists that aid in this motion. 


A muscle with the opposite action of the prime mover is called an 
antagonist. Antagonists play two important roles in muscle function: (1) 
they maintain body or limb position, such as holding the arm out or 
standing erect; and (2) they control rapid movement, as in shadow boxing 
without landing a punch or the ability to check the motion of a limb. 


For example, to extend the knee, a group of four muscles called the 
quadriceps femoris in the anterior compartment of the thigh are activated 
(and would be called the agonists of knee extension). However, to flex the 
knee joint, an opposite or antagonistic set of muscles called the hamstrings 


is activated. 


As you can see, these terms would also be reversed for the opposing action. 
If you consider the first action as the knee bending, the hamstrings would 


be called the agonists and the quadriceps femoris would then be called the 
antagonists. See [link] for a list of some agonists and antagonists. 


Agonist and Antagonist Skeletal Muscle Pairs 


Agonist 


Biceps brachii: 
in the anterior 
compartment of 
the arm 


Hamstrings: 
group of three 
muscles in the 
posterior 
compartment of 
the thigh 


Flexor 
digitorum 
superficialis 
and flexor 
digitorum 
profundus: in 
the anterior 
compartment of 
the forearm 


Antagonist 


Triceps 
brachii: in 
the posterior 
compartment 
of the arm 


Quadriceps 
femoris: 
group of 
four muscles 
in the 
anterior 
compartment 
of the thigh 


Extensor 
digitorum: in 
the posterior 
compartment 
of the 
forearm 


Movement 


The biceps brachii flexes the 
forearm, whereas the triceps 
brachii extends it. 


The hamstrings flex the leg, 
whereas the quadriceps 
femoris extend it. 


The flexor digitorum 
superficialis and flexor 
digitorum profundus flex the 
fingers and the hand at the 
wrist, whereas the extensor 
digitorum extends the fingers 
and the hand at the wrist. 


There are also skeletal muscles that do not pull against the skeleton for 
movements. For example, there are the muscles that produce facial 
expressions. The insertions and origins of facial muscles are in the skin, so 
that certain individual muscles contract to form a smile or frown, form 
sounds or words, and raise the eyebrows. There also are skeletal muscles in 
the tongue, and the external urinary and anal sphincters that allow for 
voluntary regulation of urination and defecation, respectively. In addition, 
the diaphragm contracts and relaxes to change the volume of the pleural 
cavities but it does not move the skeleton to do this. 


Note: 

Everyday Connections 

Exercise and Stretching 

When exercising, it is important to first warm up the muscles. Stretching 
pulls on the muscle fibers and it also results in an increased blood flow to 
the muscles being worked. Without a proper warm-up, it is possible that 
you may either damage some of the muscle fibers or pull a tendon. A 
pulled tendon, regardless of location, results in pain, swelling, and 
diminished function; if it is moderate to severe, the injury could 
immobilize you for an extended period. 

Recall the discussion about muscles crossing joints to create movement. 
Most of the joints you use during exercise are synovial joints, which have 
synovial fluid in the joint space between two bones. Exercise and 
stretching may also have a beneficial effect on synovial joints. Synovial 
fluid is a thin, but viscous film with the consistency of egg whites. When 
you first get up and start moving, your joints feel stiff for a number of 
reasons. After proper stretching and warm-up, the synovial fluid may 
become less viscous, allowing for better joint function. 


Patterns of Fascicle Organization 


Skeletal muscle is enclosed in connective tissue scaffolding at three levels. 
Each muscle fiber (cell) is covered by endomysium and the entire muscle is 
covered by epimysium. When a group of muscle fibers is “bundled” as a 


unit within the whole muscle by an additional covering of a connective 
tissue called perimysium, that bundled group of muscle fibers is called a 
fascicle. Fascicle arrangement by perimysia is correlated to the force 
generated by a muscle; it also affects the range of motion of the muscle. 
Based on the patterns of fascicle arrangement, skeletal muscles can be 
classified in several ways. What follows are the most common fascicle 
arrangements. 


Parallel muscles have fascicles that are arranged in the same direction as 
the long axis of the muscle ([link]). The majority of skeletal muscles in the 
body have this type of organization. Some parallel muscles are flat sheets 
that expand at the ends to make broad attachments. Other parallel muscles 
are rotund with tendons at one or both ends. Muscles that seem to be plump 
have a large mass of tissue located in the middle of the muscle, between the 
insertion and the origin, which is known as the central body. A more 
common name for this muscle is belly. When a muscle contracts, the 
contractile fibers shorten it to an even larger bulge. For example, extend and 
then flex your biceps brachii muscle; the large, middle section is the belly 
({link]). When a parallel muscle has a central, large belly that is spindle- 
shaped, meaning it tapers as it extends to its origin and insertion, it 
sometimes is called fusiform. 

Muscle Shapes and Fiber Alignment 


Orbicularis oris 


Circular 


Deltoid 


Multipennate 


Convergent 


Extensor digitorum 


Biceps brachii (posterior view) 


To origin 


Parallel- Belly 


fusiform Unipennate 


To insertion —— 


Rectus femoris 


Bipennate 


\\ 
Parallel 
( (non-fusiform) 
ma 
{ 


The skeletal muscles of the body typically come in 
seven different general shapes. 


Biceps Brachii Muscle Contraction 


The large mass at the center of a muscle is 
called the belly. Tendons emerge from 
both ends of the belly and connect the 

muscle to the bones, allowing the skeleton 

to move. The tendons of the bicep connect 
to the upper arm and the forearm. (credit: 
Victoria Garcia) 


Circular muscles are also called sphincters (see [link]). When they relax, 
the sphincters’ concentrically arranged bundles of muscle fibers increase 
the size of the opening, and when they contract, the size of the opening 
shrinks to the point of closure. The orbicularis oris muscle is a circular 
muscle that goes around the mouth. When it contracts, the oral opening 
becomes smaller, as when puckering the lips for whistling. Another 
example is the orbicularis oculi, one of which surrounds each eye. Consider, 
for example, the names of the two orbicularis muscles (orbicularis oris and 
oribicularis oculi), where part of the first name of both muscles is the same. 
The first part of orbicularis, orb (orb = “circular”), is a reference to a round 
or circular structure; it may also make one think of orbit, such as the moon’s 


path around the earth. The word oris (oris = “oral”) refers to the oral cavity, 
or the mouth. The word oculi (ocular = “eye’”) refers to the eye. 


There are other muscles throughout the body named by their shape or 
location. The deltoid is a large, triangular-shaped muscle that covers the 
shoulder. It is so-named because the Greek letter delta looks like a triangle. 
The rectus abdomis (rector = “straight”) is the straight muscle in the 
anterior wall of the abdomen, while the rectus femoris is the straight muscle 
in the anterior compartment of the thigh. 


When a muscle has a widespread expansion over a sizable area, but then the 
fascicles come to a single, common attachment point, the muscle is called 
convergent. The attachment point for a convergent muscle could be a 
tendon, an aponeurosis (a flat, broad tendon), or a raphe (a very slender 
tendon). The large muscle on the chest, the pectoralis major, is an example 
of a convergent muscle because it converges on the greater tubercle of the 
humerus via a tendon. The temporalis muscle of the cranium is another. 


Pennate muscles (penna = “feathers”) blend into a tendon that runs through 
the central region of the muscle for its whole length, somewhat like the 
quill of a feather with the muscle arranged similar to the feathers. Due to 
this design, the muscle fibers in a pennate muscle can only pull at an angle, 
and as a result, contracting pennate muscles do not move their tendons very 
far. However, because a pennate muscle generally can hold more muscle 
fibers within it, it can produce relatively more tension for its size. There are 
three subtypes of pennate muscles. 


In a unipennate muscle, the fascicles are located on one side of the tendon. 
The extensor digitorum of the forearm is an example of a unipennate 
muscle. A bipennate muscle has fascicles on both sides of the tendon. In 
some pennate muscles, the muscle fibers wrap around the tendon, 
sometimes forming individual fascicles in the process. This arrangement is 
referred to as multipennate. A common example is the deltoid muscle of 
the shoulder, which covers the shoulder but has a single tendon that inserts 
on the deltoid tuberosity of the humerus. 


Because of fascicles, a portion of a multipennate muscle like the deltoid can 
be stimulated by the nervous system to change the direction of the pull. For 


example, when the deltoid muscle contracts, the arm abducts (moves away 
from midline in the sagittal plane), but when only the anterior fascicle is 
stimulated, the arm will abduct and flex (move anteriorly at the shoulder 
joint). 


The Lever System of Muscle and Bone Interactions 


Skeletal muscles do not work by themselves. Muscles are arranged in pairs 
based on their functions. For muscles attached to the bones of the skeleton, 
the connection determines the force, speed, and range of movement. These 
characteristics depend on each other and can explain the general 
organization of the muscular and skeletal systems. 


The skeleton and muscles act together to move the body. Have you ever 
used the back of a hammer to remove a nail from wood? The handle acts as 
a lever and the head of the hammer acts as a fulcrum, the fixed point that 
the force is applied to when you pull back or push down on the handle. The 
effort applied to this system is the pulling or pushing on the handle to 
remove the nail, which is the load, or “resistance” to the movement of the 
handle in the system. Our musculoskeletal system works in a similar 
manner, with bones being stiff levers and the articular endings of the bones 
—encased in synovial joints—acting as fulcrums. The load would be an 
object being lifted or any resistance to a movement (your head is a load 
when you are lifting it), and the effort, or applied force, comes from 
contracting skeletal muscle. 


Chapter Review 


Skeletal muscles each have an origin and an insertion. The end of the 
muscle that attaches to the bone being pulled is called the muscle’s insertion 
and the end of the muscle attached to a fixed, or stabilized, bone is called 
the origin. The muscle primarily responsible for a movement is called the 
prime mover, and muscles that assist in this action are called synergists. A 
synergist that makes the insertion site more stable is called a fixator. 
Meanwhile, a muscle with the opposite action of the prime mover is called 
an antagonist. Several factors contribute to the force generated by a skeletal 
muscle. One is the arrangement of the fascicles in the skeletal muscle. 


Fascicles can be parallel, circular, convergent, pennate, fusiform, or 
triangular. Each arrangement has its own range of motion and ability to do 
work. 


Review Questions 


Exercise: 


Problem: 
Which of the following is unique to the muscles of facial expression? 


a. They all originate from the scalp musculature. 

b. They insert onto the cartilage found around the face. 
c. They only insert onto the facial bones. 

d. They insert into the skin. 


Solution: 


D 


Exercise: 


Problem: Which of the following helps an agonist work? 


a. a Synergist 
b. a fixator 

c. an insertion 
d. an antagonist 


Solution: 


A 


Exercise: 


Problem: 


Which of the following statements is correct about what happens 
during flexion? 


a. The angle between bones is increased. 

b. The angle between bones is decreased. 

c. The bone moves away from the body. 

d. The bone moves toward the center of the body. 


Solution: 


B 


Exercise: 


Problem: Which is moved the /east during muscle contraction? 


a. the origin 
b. the insertion 
c. the ligaments 
d. the joints 


Solution: 


A 


Exercise: 


Problem: Which muscle has a convergent pattern of fascicles? 


a. biceps brachii 
b. gluteus maximus 
c. pectoralis major 
d. rectus femoris 


Solution: 


C 
Exercise: 


Problem: 


A muscle that has a pattern of fascicles running along the long axis of 
the muscle has which of the following fascicle arrangements? 


a. Circular 
b. pennate 
c. parallel 
d. rectus 


Solution: 


C 


Exercise: 


Problem: Which arrangement best describes a bipennate muscle? 


a. The muscle fibers feed in on an angle to a long tendon from both 
sides. 

b. The muscle fibers feed in on an angle to a long tendon from all 
directions. 

c. The muscle fibers feed in on an angle to a long tendon from one 
side. 

d. The muscle fibers on one side of a tendon feed into it at a certain 
angle and muscle fibers on the other side of the tendon feed into it 
at the opposite angle. 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: 
What effect does fascicle arrangement have on a muscle’s action? 
Solution: 


Fascicle arrangements determine what type of movement a muscle can 

make. For instance, circular muscles act as sphincters, closing orifices. 
Exercise: 

Problem: 


Movements of the body occur at joints. Describe how muscles are 
arranged around the joints of the body. 


Solution: 


Muscles work in pairs to facilitate movement of the bones around the 
joints. Agonists are the prime movers while antagonists oppose or 
resist the movements of the agonists. Synergists assist the agonists, and 
fixators stabilize a muscle’s origin. 


Exercise: 
Problem:Explain how a synergist assists an agonist by being a fixator. 
Solution: 
Agonists are the prime movers while antagonists oppose or resist the 


movements of the agonists. Synergists assist the agonists, and fixators 
stabilize a muscle’s origin. 


Glossary 


abduct 
move away from midline in the sagittal plane 


agonist 
(also, prime mover) muscle whose contraction is responsible for 
producing a particular motion 


antagonist 
muscle that opposes the action of an agonist 


belly 
bulky central body of a muscle 


bipennate 
pennate muscle that has fascicles that are located on both sides of the 
tendon 


circular 
(also, sphincter) fascicles that are concentrically arranged around an 
opening 


convergent 
fascicles that extend over a broad area and converge on a common 
attachment site 


fascicle 
muscle fibers bundled by perimysium into a unit 


fixator 
synergist that assists an agonist by preventing or reducing movement at 
another joint, thereby stabilizing the origin of the agonist 


flexion 
movement that decreases the angle of a joint 


fusiform 
muscle that has fascicles that are spindle-shaped to create large bellies 


insertion 


end of a skeletal muscle that is attached to the structure (usually a 
bone) that is moved when the muscle contracts 


multipennate 
pennate muscle that has a tendon branching within it 


origin 
end of a skeletal muscle that is attached to another structure (usually a 
bone) in a fixed position 


parallel 
fascicles that extend in the same direction as the long axis of the 
muscle 


pennate 
fascicles that are arranged differently based on their angles to the 
tendon 


prime mover 
(also, agonist) principle muscle involved in an action 


synergist 
muscle whose contraction helps a prime mover in an action 


unipennate 
pennate muscle that has fascicles located on one side of the tendon 


Naming Skeletal Muscles 
By the end of this section, you will be able to: 


e Describe the criteria used to name skeletal muscles 
e Explain how understanding the muscle names helps describe shapes, 
location, and actions of various muscles 


The Greeks and Romans conducted the first studies done on the human 
body in Western culture. The educated class of subsequent societies studied 
Latin and Greek, and therefore the early pioneers of anatomy continued to 
apply Latin and Greek terminology or roots when they named the skeletal 
muscles. The large number of muscles in the body and unfamiliar words 
can make learning the names of the muscles in the body seem daunting, but 
understanding the etymology can help. Etymology is the study of how the 
root of a particular word entered a language and how the use of the word 
evolved over time. Taking the time to learn the root of the words is crucial 
to understanding the vocabulary of anatomy and physiology. When you 
understand the names of muscles it will help you remember where the 
muscles are located and what they do ({link], [link], and [link]). 
Pronunciation of words and terms will take a bit of time to master, but after 
you have some basic information; the correct names and pronunciations 
will become easier. 

Overview of the Muscular System 


‘ Occipitofrontalis 
Sternocleidomastoid i ———— (frontal belly) 


Deltoid 
Trapezius 
Pectoralis major Pectoralis minor 


ae Serratus anterior 
Rectus abdominis 


Biceps brachii 
Abdominal Brachialis 


external oblique 


Brachioradialis 


: Pronator teres 
Pectineus 


Flexor carpi radialis 
Adductor Tensor fasciae latae 
longus 


Sartorius 


llliopsoas 
Rectus femoris Js 
Gracilis 


Vastus lateralis Vastus medialis 
Soleus and 


Fibularis longus : 
gastrocnemius 


Tibialis anterior 


a) ip 


Major muscles of the body. 
Right side: superficial; left side: 
deep (anterior view) 


Occipitofrontalis 


(occipital belly) Epicranial aponeurosis 


Splenius capitis 
Levator scapulae 


Supraspinatus Rhomboids 
i Trapezius 
Teres minor 
Deltoid 


Infraspinatus 
f Latissimus dorsi 
Teres major 
: ri Brachioradialis 
Triceps brachii ee 
: Extensor carpi radialis 
Serratus posterior 


inferior Extensor digitorum 


External oblique Extensor carpi ulnaris 


Gluteus medius Flexor carpi ulnaris 
(dissected) 
Gluteus maximus 77] 
(dissected) 


Semimembranosus 


“> Gluteus minimus 
i \ + Gemellus muscles 
“YN 


Biceps femoris 
Semitendinosus 

Gracilis 

Gastrocnemius (dissected) 


Peroneus longus 


Tibialis posterior Soleus 


Major muscles of the body. 
Right side: superficial; left side: 
deep (posterior view) 


On the anterior and posterior views of the 
muscular system above, superficial 
muscles (those at the surface) are shown 
on the right side of the body while deep 
muscles (those underneath the superficial 
muscles) are shown on the left half of the 
body. For the legs, superficial muscles are 
shown in the anterior view while the 
posterior view shows both superficial and 
deep muscles. 


Understanding a Muscle Name from the Latin 


abductor ab = away from duct = to move a Ieee 
moves away from 
A muscle that 
abductor F 
diciti diaiti digitus = digit refers to a finger moves the 
gmt 9 9 9 or toe little finger or 
minimi 
oh: toe away 
pa minimus = : 
minimi ase little 
mini, tiny 


adductor ad = to, toward duct = to move enindscle te 
moves towards 
Sdduictor A muscle that 
diciti diaiti diaitus = diait refers to a finger moves the 
ont g 9 9 or toe little finger or 
minimi 
aE toe toward 
8 cent minimus = \ 
minimi ae little 
mini, tiny 


Mnemonic Device for Latin Roots 


Latin or 
Greek 
Example Translation Mnemonic Device 
ad to; toward ADvance toward your goal 
ab away from n/a 
sub under SUBmarines move under water. 
something A conDUCTOR makes a train 
ductor 
that moves move. 
If you are antisocial, you are 
anti against against engaging in social 
activities. 
epi on top of n/a 


apo to the side of n/a 


Mnemonic Device for Latin Roots 


Example 


longissimus 


longus 
brevis 


maximus 


medius 


minimus 


rectus 


multi 


uni 


bi/di 


tri 


quad 


Latin or 
Greek 


Translation 


longest 


long 
short 


large 


medium 


tiny; little 


straight 


many 


one 


two 


three 


four 


Mnemonic Device 


“Longissimus” is longer than the 
word “long.” 


long 
brief 
max 


“Medius” and “medium” both 
begin with “med.” 


mini 


To RECTify a situation is to 
straighten it out. 


If something is MULTIcolored, it 
has many colors. 


A UNIcorn has one horn. 


If a ring is DIcast, it is made of 
two metals. 


TRIple the amount of money is 
three times as much. 


QUADruplets are four children 
born at one birth. 


Mnemonic Device for Latin Roots 


Latin or 

Greek 
Example Translation Mnemonic Device 
externus outside EXternal 
internus inside INternal 


Anatomists name the skeletal muscles according to a number of criteria, 
each of which describes the muscle in some way. These include naming the 
muscle after its shape, its size compared to other muscles in the area, its 
location in the body or the location of its attachments to the skeleton, how 
many origins it has, or its action. 


The skeletal muscle’s anatomical location or its relationship to a particular 
bone often determines its name. For example, the frontalis muscle is located 
on top of the frontal bone of the skull. Similarly, the shapes of some 
muscles are very distinctive and the names, such as orbicularis, reflect the 
shape. For the buttocks, the size of the muscles influences the names: 
gluteus maximus (largest), gluteus medius (medium), and the gluteus 
minimus (smallest). Names were given to indicate length—brevis (short), 
longus (long)—and to identify position relative to the midline: lateralis (to 
the outside away from the midline), and medialis (toward the midline). The 
direction of the muscle fibers and fascicles are used to describe muscles 
relative to the midline, such as the rectus (straight) abdominis, or the 
oblique (at an angle) muscles of the abdomen. 


Some muscle names indicate the number of muscles in a group. One 
example of this is the quadriceps, a group of four muscles located on the 
anterior (front) thigh. Other muscle names can provide information as to 
how many origins a particular muscle has, such as the biceps brachii. The 
prefix bi indicates that the muscle has two origins and tri indicates three 
origins. 


The location of a muscle’s attachment can also appear in its name. When 
the name of a muscle is based on the attachments, the origin is always 
named first. For instance, the sternocleidomastoid muscle of the neck has a 
dual origin on the sternum (sterno) and clavicle (cleido), and it inserts on 
the mastoid process of the temporal bone. The last feature by which to 
name a muscle is its action. When muscles are named for the movement 
they produce, one can find action words in their name. Some examples are 
flexor (decreases the angle at the joint), extensor (increases the angle at the 
joint), abductor (moves the bone away from the midline), or adductor 
(moves the bone toward the midline). 


Chapter Review 


Muscle names are based on many characteristics. The location of a muscle 
in the body is important. Some muscles are named based on their size and 
location, such as the gluteal muscles of the buttocks. Other muscle names 
can indicate the location in the body or bones with which the muscle is 
associated, such as the tibialis anterior. The shapes of some muscles are 
distinctive; for example, the direction of the muscle fibers is used to 
describe muscles of the body midline. The origin and/or insertion can also 
be features used to name a muscle; examples are the biceps brachii, triceps 
brachii, and the pectoralis major. 


Review Questions 


Exercise: 


Problem: 


The location of a muscle’s insertion and origin can determine 


a. action 

b. the force of contraction 

c. muscle name 

d. the load a muscle can carry 


Solution: 


A 


Exercise: 


Problem: Where is the temporalis muscle located? 


a. on the forehead 

b. in the neck 

c. on the side of the head 
d. on the chin 


Solution: 


c 


Exercise: 


Problem:Which muscle name does not make sense? 


a. extensor digitorum 

b. gluteus minimus 

c. biceps femoris 

d. extensor minimus longus 


Solution: 


D 
Exercise: 


Problem: 


Which of the following terms would be used in the name of a muscle 
that moves the leg away from the body? 


a. flexor 


b. adductor 
c. extensor 
d. abductor 


Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: 


Describe the different criteria that contribute to how skeletal muscles 
are named. 


Solution: 


In anatomy and physiology, many word roots are Latin or Greek. 
Portions, or roots, of the word give us clues about the function, shape, 
action, or location of a muscle. 


Glossary 


abductor 
moves the bone away from the midline 


adductor 
moves the bone toward the midline 


bi 
two 


brevis 
short 


extensor 
muscle that increases the angle at the joint 


flexor 
muscle that decreases the angle at the joint 


lateralis 
to the outside 


longus 
long 


maximus 
largest 


medialis 
to the inside 


medius 
medium 


minimus 
smallest 


oblique 
at an angle 


rectus 
straight 


tri 
three 


Muscles of the Head, Neck, and Back 
By the end of this section, you will be able to: 


e Identify the axial muscles of the face, head, and neck 
e Identify the movement and function of the face, head, and neck muscles 


The skeletal muscles are divided into axial (muscles of the trunk and head) and appendicular (muscles of 
the arms and legs) categories. This system reflects the bones of the skeleton system, which are also 
arranged in this manner. The axial muscles are grouped based on location, function, or both. Some of the 
axial muscles may seem to blur the boundaries because they cross over to the appendicular skeleton. The 
first grouping of the axial muscles you will review includes the muscles of the head and neck, then you 
will review the muscles of the vertebral column, and finally you will review the oblique and rectus 
muscles. 


Muscles That Create Facial Expression 


The origins of the muscles of facial expression are on the surface of the skull (remember, the origin of a 
muscle does not move). The insertions of these muscles have fibers intertwined with connective tissue and 
the dermis of the skin. Because the muscles insert in the skin rather than on bone, when they contract, the 
skin moves to create facial expression ([link]). 

Muscles of Facial Expression 


Epicranial aponeurosis 


Occipitofrontalis 
(frontal belly) 


Corrugator supercilii 


Orbicularis oculi 


Occipitofrontalis 
(occipital belly) 


Orbicularis oris 


Facial muscles (anterior view) Facial muscles (lateral view) 


Many of the muscles of facial expression insert into the 
skin surrounding the eyelids, nose and mouth, producing 
facial expressions by moving the skin rather than bones. 


The orbicularis oris is a circular muscle that moves the lips, and the orbicularis oculi is a circular muscle 
that closes the eye. The occipitofrontalis muscle moves up the scalp and eyebrows. The muscle has a 
frontal belly and an occipital (near the occipital bone on the posterior part of the skull) belly. In other 
words, there is a muscle on the forehead (frontalis) and one on the back of the head (occipitalis), but there 
is no muscle across the top of the head. Instead, the two bellies are connected by a broad tendon called the 
epicranial aponeurosis, or galea aponeurosis (galea = “apple”). The physicians originally studying human 
anatomy thought the skull looked like an apple. 


A large portion of the face is composed of the buccinator muscle, which compresses the cheek. This 
muscle allows you to whistle, blow, and suck; and it contributes to the action of chewing. There are 


several small facial muscles, one of which is the corrugator supercilii, which is the prime mover of the 
eyebrows. Place your finger on your eyebrows at the point of the bridge of the nose. Raise your eyebrows 
as if you were surprised and lower your eyebrows as if you were frowning. With these movements, you 
can feel the action of the corrugator supercilli. Additional muscles of facial expression are presented in 
[link]. 

Muscles in Facial Expression 


Target motion 
direction 


Brow 

Furrowing Skin of scalp Anterior Occipito- Epicraneal Underneath 

brow frontalis, aponeurosis skin of 
frontal belly forehead 

Unfurrowing Skin of scalp Posterior Occipito- Occipital bone; Epicraneal 

brow frontalis, mastoid process aponeurosis 
occipital belly (temporal bone) 

Lowering Skin Inferior Corrugator Frontal bone Skin 

eyebrows underneath supercilii underneath 

(e.g., scowling, eyebrows eyebrow 

frowning) 


Nose 


Movement Target Prime mover Origin Insertion 


Flaring nostrils Nasal cartilage Inferior Nasal bone 
(pushes nostrils | compression; 
open when posterior 
cartilage is compression 
compressed) 


Raising Upper lip Elevation Levator labii Maxilla Underneath 
upper lip superioris skin at corners 
of the mouth; 
orbicularis oris 


Lowering Lower lip Depression Depressor Mandible Underneath 
lower lip labii inferioris skin of lower lip 


Opening mouth | Lower jaw Depression, Depressor Mandible Underneath 
and sliding lateral angulus oris skin at corners 
lower jaw left of mouth 

and right 

Smiling Corners of Lateral Zygomaticus Zygomatic bone Underneath 


skin at corners 
of mouth 

(dimple area); 
orbicularis oris 


Shaping of lips Lips Orbicularis Tissue Underneath 
(as during oris surrounding lips skin at corners 
speech) of the mouth 


Lateral Cheeks Lateral Buccinator Maxilla, mandible; | Orbicularis 
movement of sphenoid bone (via | oris 
cheeks (e.g., pterygomandibular 

sucking on a raphae) 

straw; also used 

to compress air 

in mouth while 

blowing) 


Pursing of lips Corners of Lateral Risorius Fascia of parotid Underneath 
by straightening | mouth salivary gland skin at corners 
them laterally of the mouth 


Mandible Underneath 
skin of chin 


mouth elevation major 


Protrusion of Lower lip and Protraction Mentalis 
lower lip (e.g., skin of chin 

pouting 

expression) 


Muscles That Move the Eyes 


The movement of the eyeball is under the control of the extrinsic eye muscles, which originate outside the 
eye and insert onto the outer surface of the white of the eye. These muscles are located inside the eye 
socket and cannot be seen on any part of the visible eyeball ({link] and [link]). If you have ever been to a 
doctor who held up a finger and asked you to follow it up, down, and to both sides, he or she is checking 
to make sure your eye muscles are acting in a coordinated pattern. 

Muscles of the Eyes 


Superior oblique 


Levator palpebrae superioris 


Superior oblique 


Superior 4 
rectus ] Lateral 


rectus 


Sphenoid 
bone 


~ eS 
S33 » Medial 
: oe ge % 
we Lateral \ ey Ee “= Ye, rectus 
i rectus Ss aeet 
Inferior rectus & 3 
ge 
Sa = 
Medial rectus Inferior oblique Inferior oblique Inferior rectus 
(a) Right eye (lateral view) (b) Right eye (anterior view) 


(a) The extrinsic eye muscles originate outside of the eye on 
the skull. (b) Each muscle inserts onto the eyeball. 


Muscles of the Eyes 


Target 

motion Prime 
Movement Target direction mover 
Moves eyes up 
and toward Superior 
nose; rotates Eyeballs (elevates); Superior 
eyes from 1 medial rectus 
o’clock to 3 (adducts) 
o’clock 
Moves eyes 
down and Inferior 
toward nose; Eyeballs (depresses); Inferior 
rotates eyes medial rectus 
from 6 o’clock (adducts) 
to 3 o’clock 
mea Lateral Lateral 
away from Eyeballs 
‘gee (abducts) rectus 


Origin 


Common 
tendinous 
ring (ring 
attaches to 
optic 
foramen) 


Common 
tendinous 
ring (ring 
attaches to 
optic 
foramen) 


Common 
tendinous 
ring (ring 
attaches to 
optic 
foramen) 


Insertion 


Superior 
surface of 
eyeball 


Inferior 
surface of 
eyeball 


Lateral 
surface of 
eyeball 


Muscles of the Eyes 


Movement 


Moves eyes 
toward nose 


Moves eyes up 
and away from 
nose; rotates 
eyeball from 
12 o’clock to 9 
o’clock 


Moves eyes 
down and 
away from 
nose; rotates 
eyeball from 6 
o’clock to 9 
o’clock 


Opens eyes 


Closes eyelids 


Target 


Eyeballs 


Eyeballs 


Eyeballs 


Upper 
eyelid 


Eyelid 
skin 


Target 
motion 
direction 


Medial 
(adducts) 


Superior 
(elevates); 
lateral 
(abducts) 


Superior 
(elevates); 
lateral 
(abducts) 


Superior 
(elevates) 


Compression 
along 
superior— 
inferior axis 


Muscles That Move the Lower Jaw 


Prime 
mover 


Medial 
rectus 


Inferior 
oblique 


Superior 
oblique 


Levator 
palpabrae 
superioris 


Orbicularis 
oculi 


Origin 


Common 
tendinous 
ring (ring 
attaches to 
optic 
foramen) 


Floor of 
orbit 
(maxilla) 


Sphenoid 
bone 


Roof of 
orbit 
(sphenoid 
bone) 


Medial 
bones 
composing 
the orbit 


Insertion 


Medial 
surface of 
eyeball 


Surface of 
eyeball 
between 
inferior rectus 
and lateral 
rectus 


Suface of 
eyeball 
between 
superior rectus 
and lateral 
rectus 


Skin of upper 
eyelids 


Circumference 
of orbit 


In anatomical terminology, chewing is called mastication. Muscles involved in chewing must be able to 
exert enough pressure to bite through and then chew food before it is swallowed ([link] and [link]). The 
masseter muscle is the main muscle used for chewing because it elevates the mandible (lower jaw) to 
close the mouth, and it is assisted by the temporalis muscle, which retracts the mandible. You can feel the 


temporalis move by putting your fingers to your temple as you chew. 
Muscles That Move the Lower Jaw 


Lateral 
pterygoid 


Area of 
superficial 
muscle 

dissection 


Medial 
pterygoid 


Chewing muscles (superficial) Chewing muscles (deep) 


The muscles that move the lower jaw are typically 
located within the cheek and originate from processes 
in the skull. This provides the jaw muscles with the 
large amount of leverage needed for chewing. 


Muscles of the Lower Jaw 


Target 
motion Prime 
Movement Target direction mover Origin Insertion 
Maxilla 
. : arch; 
ea ou Mandible supenor Masseter zygomatic Mandible 
aids chewing (elevates) 
arch (for 
masseter) 
Closes mouth; Superior 
pulls Lower jaw Mandible (elevates); Temporalis Jemporal Mandible 
in under upper posterior bone 
jaw (retracts) 
Inferior 
Opens mouth; (depresses); 
pushes lower posterior Pterygoid 
jaw out under Mandible (protracts); Lateral process of Mandible 
upper jaw; lateral pterygoid sphenoid 
moves lower (abducts); bone 
jaw side-to-side medial 


(adducts) 


Muscles of the Lower Jaw 


Target 
motion Prime 
Movement Target direction mover Origin Insertion 
Superior 
Closes mouth; (elevates); 
pushes lower posterior . Mandible; 
: . Sphenoid 
jaw out under : (protracts); Medial : temporo- 
: Mandible : bone; : 
upper jaw; lateral pterygoid faawilla mandibular 
moves lower (abducts); joint 
jaw side-to-side medial 
(adducts) 


Although the masseter and temporalis are responsible for elevating and closing the jaw to break food into 
digestible pieces, the medial pterygoid and lateral pterygoid muscles provide assistance in chewing and 
moving food within the mouth. 


Muscles That Move the Tongue 


Although the tongue is obviously important for tasting food, it is also necessary for mastication, 
deglutition (swallowing), and speech ({link] and [link]). Because it is so moveable, the tongue facilitates 
complex speech patterns and sounds. 

Muscles that Move the Tongue 


Styloglossus ~~ 


Pharyngopalatine arch 


Dorsal surface 


Palatine tonsil 
of tongue 


Palatoglossus 


Buccinator 
Hyoglossus 


i Fungiform 
Valate Z ; i papilla 

papilla Fe 
Mandible bone 


Genioglossus 


(a) Extrinsic tongue muscles (b) Palatoglossus and surface of tongue 


Muscles for Tongue Movement, Swallowing, and Speech 


Target motion Prime 
ene ae | Tgecnaren | Rome | onan | insertion | 


Tongue 
Moves tongue down; sticks Tongue Inferior (depresses); | Genioglossus | Mandible Tongue 
tongue out of mouth anterior (protracts) undersurface; 
hyoid bone 
Moves tongue up; retracts Tongue Superior (elevates); | Styloglossus Temporal Tongue 
tongue back into mouth posterior (retracts) bone (styloid | undersurface 
process) and sides 


Flattens tongue Tongue Inferior (depresses) | Hyoglossus Hyoid bone Sides of 
tongue 

Bulges tongue Tongue Superior (elevation) | Palatoglossus | Soft palate 
tongue 


Swallowing and speaking 


Raises the hyoid bone in a way Hyoid bone; | Superior (elevates) Digastric Mandible; Hyoid bone 
that also raises the larynx, larynx temporal 

allowing the epiglottis to cover bone 

the glottis during deglutition; 

also assists in opening the 

mouth by depressing the 

mandible 


Raises and retracts the hyoid Hyoid bone | Superior (elevates); | Stylohyoid Temporal Hyoid bone 
bone in a way that elongates posterior (retracts) bone (styloid 
the oral cavity during deglutition process) 


Mylohyoid Mandible Hyoid bone; 


Raises hyoid bone in a way Hyoid bone | Superior (elevates) 
that presses tongue against 
the roof of the mouth, pushing 
food back into the pharynx 
during deglutition 


Raises and moves hyoid bone Hyoid bone | Superior (elevates); | Geniohyoid Mandible Hyoid bone 
forward, widening pharynx anterior (protracts) 

during deglutition 

Retracts hyoid bone and Hyoid bone | Inferior (depresses); | Omohyoid Scapula Hyoid bone 
moves it down during later posterior (retracts) 

phases of deglutition 

Depresses the hyoid bone Hyoid bone | Inferior (depresses) | Sternohyoid Clavicle Hyoid bone 
during swallowing and speaking 


Shrinks distance between Hyoid bone; | Hyoid bone: inferior | Thyrohyoid Thyroid Hyoid bone 
thyroid cartilage and hyoid thyroid (depresses); thyroid cartilage 

bone, allowing production of cartilage cartilage: superior 

high-pitch vocalizations (elevates) 


Depresses larynx, thyroid Larynx; Inferior (depresses) | Sternothyroid | Sternum Thyroid 
cartilage, and hyoid bone to thyroid cartilage 
create different vocal tones cartilage; 

hyoid bone 


median raphe 


Rotates and tilts head to he Skull; Individually: medial Sternocleid- Sternum; Temporal bone 
side; tilts head forward cervical rotation; lateral omastoid; clavicle (mastoid 
vertebrae flexion; bilaterally: semispinalis Process); 
anterior (flexes) capitis occipital bone 
Rotates and tilts head to the Skull; Individually: lateral Splenius 
side; tilts head backwards cervical rotation; lateral capitis; 
vertebrae flexion; bilaterally: longissimus 
anterior (flexes) capitis 


Tongue muscles can be extrinsic or intrinsic. Extrinsic tongue muscles insert into the tongue from outside 
origins, and the intrinsic tongue muscles insert into the tongue from origins within it. The extrinsic 
muscles move the whole tongue in different directions, whereas the intrinsic muscles allow the tongue to 
change its shape (such as, curling the tongue in a loop or flattening it). 


The extrinsic muscles all include the word root glossus (glossus = “tongue’”’), and the muscle names are 
derived from where the muscle originates. The genioglossus (genio = “chin”) originates on the mandible 
and allows the tongue to move downward and forward. The styloglossus originates on the styloid bone, 
and allows upward and backward motion. The palatoglossus originates on the soft palate to elevate the 
back of the tongue, and the hyoglossus originates on the hyoid bone to move the tongue downward and 
flatten it. 


Note: 

Everyday Connections 

Anesthesia and the Tongue Muscles 

Before surgery, a patient must be made ready for general anesthesia. The normal homeostatic controls of 
the body are put “on hold” so that the patient can be prepped for surgery. Control of respiration must be 
switched from the patient’s homeostatic control to the control of the anesthesiologist. The drugs used for 
anesthesia relax a majority of the body’s muscles. 


Among the muscles affected during general anesthesia are those that are necessary for breathing and 
moving the tongue. Under anesthesia, the tongue can relax and partially or fully block the airway, and the 
muscles of respiration may not move the diaphragm or chest wall. To avoid possible complications, the 
safest procedure to use on a patient is called endotracheal intubation. Placing a tube into the trachea 
allows the doctors to maintain a patient’s (open) airway to the lungs and seal the airway off from the 
oropharynx. Post-surgery, the anesthesiologist gradually changes the mixture of the gases that keep the 
patient unconscious, and when the muscles of respiration begin to function, the tube is removed. It still 
takes about 30 minutes for a patient to wake up, and for breathing muscles to regain control of respiration. 
After surgery, most people have a sore or scratchy throat for a few days. 


Muscles of the Anterior Neck 


The muscles of the anterior neck assist in deglutition (swallowing) and speech by controlling the positions 
of the larynx (voice box), and the hyoid bone, a horseshoe-shaped bone that functions as a solid 
foundation on which the tongue can move. The muscles of the neck are categorized according to their 
position relative to the hyoid bone ([{link]). Suprahyoid muscles are superior to it, and the infrahyoid 
muscles are located inferiorly. 

Muscles of the Anterior Neck 


Suprahyoid 
muscles: 


Geniohyoid 
Digastric 
Mylohyoid 
Stylohyoid 


Inferior edge 
of mandible 


Styloglossus 
Hyoid bone 


Infrahyoid 
muscles: 


Thyroid cartilage 
Thyrohyoid 


of larynx 
Omohyoid 


Thyroid gland J Sternohyoid 
Sternothyroid 


Trachea 
Right and left 


—e_cclavicles 


L— ~ Sternum 


The anterior muscles of the neck facilitate 
swallowing and speech. The suprahyoid 
muscles originate from above the hyoid bone 
in the chin region. The infrahyoid muscles 
originate below the hyoid bone in the lower 
neck. 


Scapula 


The suprahyoid muscles raise the hyoid bone, the floor of the mouth, and the larynx during deglutition. 
These include the digastric muscle, which has anterior and posterior bellies that work to elevate the hyoid 
bone and larynx when one swallows; it also depresses the mandible. The stylohyoid muscle moves the 
hyoid bone posteriorly, elevating the larynx, and the mylohyoid muscle lifts it and helps press the tongue 
to the top of the mouth. The geniohyoid depresses the mandible in addition to raising and pulling the 
hyoid bone anteriorly. 


The strap-like infrahyoid muscles generally depress the hyoid bone and control the position of the larynx. 
The omohyoid muscle, which has superior and inferior bellies, depresses the hyoid bone in conjunction 
with the sternohyoid and thyrohyoid muscles. The thyrohyoid muscle also elevates the larynx’s thyroid 
cartilage, whereas the sternothyroid depresses it to create different tones of voice. 


Muscles That Move the Head 


The head, attached to the top of the vertebral column, is balanced, moved, and rotated by the neck muscles 
([link]). When these muscles act unilaterally, the head rotates. When they contract bilaterally, the head 
flexes or extends. The major muscle that laterally flexes and rotates the head is the sternocleidomastoid. 
In addition, both muscles working together are the flexors of the head. Place your fingers on both sides of 
the neck and turn your head to the left and to the right. You will feel the movement originate there. This 
muscle divides the neck into anterior and posterior triangles when viewed from the side ([link]). 
Posterior and Lateral Views of the Neck 

(> = = ; Suboccipital muscles 


= < ” Splenius 
a \S capitis (cut) 


Sternocleidomastoid Levator 


Levator scapulae 

| . Longissimus 
Multifidus . i capitis 
muscles ‘ \ 


Acromion 
process of Semispinalis 
capitis 


1st thoracic 
vertebrae 


Sma Scalenes 


Neck muscles Superficial neck muscles: Deep neck muscles: left 
(left lateral view) tight side trapezius removed side semispinalis capitis 
(posterior view) removed (posterior view) 


The superficial and deep muscles of the neck are responsible for moving the 
head, cervical vertebrae, and scapulas. 


Muscles That Move the Head 


Target 

motion 
Movement Target direction Prime mover Origin Insertion 
Rotates Individually: Temporal 
and tilts rotates head bone 
head to the Skull; to opposite Grcmuleidemacuid Sternum; (mastoid 
side; tilts vertebrae side; clavicle process); 
head bilaterally: occipital 


forward flexion bone 


Muscles That Move the Head 


Target 

motion 
Movement Target direction Prime mover Origin Insertion 

Individually: Transverse 

laterally and 
Rotates flexes and articular 
and tilts Skull; rotates head Semispinalis capitis processes Occipital 
head vertebrae to same of cervical bone 
backward side; and 

bilaterally: thoracic 

extension vertebra 

Individually: 
Rotates laterally Spinous Temporal 
and tilts flexes and processes bone 
head to the Skull; rotates head Sian niccanits of cervical (mastoid 
side; tilts vertebrae to same P P and process); 
head side; thoracic occipital 
backward bilaterally: vertebra bone 

extension 

Individually: Transverse 
Rotates laterally and 
and tilts flexes and articular Temporal 
head to the Skull; rotates head Longissimus capitis processes bone 
side; tilts vertebrae to same of cervical (mastoid 
head side; and process) 
backward bilaterally: thoracic 

extension vertebra 


Muscles of the Posterior Neck and the Back 


The posterior muscles of the neck are primarily concerned with head movements, like extension. The back 
muscles stabilize and move the vertebral column, and are grouped according to the lengths and direction 


of the fascicles. 


The splenius muscles originate at the midline and run laterally and superiorly to their insertions. From the 
sides and the back of the neck, the splenius capitis inserts onto the head region, and the splenius cervicis 
extends onto the cervical region. These muscles can extend the head, laterally flex it, and rotate it ([link]). 
Muscles of the Neck and Back 


Sternocleidomastoid 


Trapezius 


Splenius capitis 


Splenius 


Splenius cervicis 
Levator 


scapulae P 
Rhomboides 


minor 


Rhomboides 
major 


Trapezius 
Medial 
scalene 


Anterior 
scalene 


Muscles of the neck (left lateral view) Superficial (left side) and deep 
(right side) muscles of the neck and 
upper back (posterior view) 


Semispinalis capitis 4 Longissimus capitis 
(joined with deep 
spinalis capitis) ql lliocostalis cervicis 


Semispinalis S lliocostalis thoracis 
cervicis , 


ee Longissimus thoracis 
Longissimus 

cervicis 
lliocostalis lumborum 


Spinalis 
thoracis Transverse 
processes 


of vertebrae 


Semispinalis i / Rotator 
thoracis H brevis 


Rotator 
longus 


Interspinales 


Short 
rotator 


Intertransversarii 


Deep muscles of the back Deep spinal muscles 
(posterior view) (multifidus removed) 


The large, complex muscles of the neck and back move the 
head, shoulders, and vertebral column. 


The erector spinae group forms the majority of the muscle mass of the back and it is the primary extensor 
of the vertebral column. It controls flexion, lateral flexion, and rotation of the vertebral column, and 
maintains the lumbar curve. The erector spinae comprises the iliocostalis (laterally placed) group, the 
longissimus (intermediately placed) group, and the spinalis (medially placed) group. 


The iliocostalis group includes the iliocostalis cervicis, associated with the cervical region; the 
iliocostalis thoracis, associated with the thoracic region; and the iliocostalis lumborum, associated with 
the lumbar region. The three muscles of the longissimus group are the longissimus capitis, associated 
with the head region; the longissimus cervicis, associated with the cervical region; and the longissimus 
thoracis, associated with the thoracic region. The third group, the spinalis group, comprises the spinalis 
capitis (head region), the spinalis cervicis (cervical region), and the spinalis thoracis (thoracic region). 


The transversospinales muscles run from the transverse processes to the spinous processes of the 
vertebrae. Similar to the erector spinae muscles, the semispinalis muscles in this group are named for the 
areas of the body with which they are associated. The semispinalis muscles include the semispinalis 
capitis, the semispinalis cervicis, and the semispinalis thoracis. The multifidus muscle of the lumbar 
region helps extend and laterally flex the vertebral column. 


Important in the stabilization of the vertebral column is the segmental muscle group, which includes the 
interspinales and intertransversarii muscles. These muscles bring together the spinous and transverse 


processes of each consecutive vertebra. Finally, the scalene muscles work together to flex, laterally flex, 
and rotate the head. They also contribute to deep inhalation. The scalene muscles include the anterior 
scalene muscle (anterior to the middle scalene), the middle scalene muscle (the longest, intermediate 
between the anterior and posterior scalenes), and the posterior scalene muscle (the smallest, posterior to 
the middle scalene). 


Chapter Review 


Muscles are either axial muscles or appendicular. The axial muscles are grouped based on location, 
function, or both. Some axial muscles cross over to the appendicular skeleton. The muscles of the head 
and neck are all axial. The muscles in the face create facial expression by inserting into the skin rather than 
onto bone. Muscles that move the eyeballs are extrinsic, meaning they originate outside of the eye and 
insert onto it. Tongue muscles are both extrinsic and intrinsic. The genioglossus depresses the tongue and 
moves it anteriorly; the styloglossus lifts the tongue and retracts it; the palatoglossus elevates the back of 
the tongue; and the hyoglossus depresses and flattens it. The muscles of the anterior neck facilitate 
swallowing and speech, stabilize the hyoid bone and position the larynx. The muscles of the neck stabilize 
and move the head. The sternocleidomastoid divides the neck into anterior and posterior triangles. 


The muscles of the back and neck that move the vertebral column are complex, overlapping, and can be 
divided into five groups. The splenius group includes the splenius capitis and the splenius cervicis. The 
erector spinae has three subgroups. The iliocostalis group includes the iliocostalis cervicis, the iliocostalis 
thoracis, and the iliocostalis lumborum. The longissimus group includes the longissimus capitis, the 
longissimus cervicis, and the longissimus thoracis. The spinalis group includes the spinalis capitis, the 
spinalis cervicis, and the spinalis thoracis. The transversospinales include the semispinalis capitis, 
semispinalis cervicis, semispinalis thoracis, multifidus, and rotatores. The segmental muscles include the 
interspinales and intertransversarii. Finally, the scalenes include the anterior scalene, middle scalene, and 
posterior scalene. 


Review Questions 


Exercise: 


Problem: Which of the following is a prime mover in head flexion? 


a. occipitofrontalis 

b. corrugator supercilii 
c. sternocleidomastoid 
d. masseter 


Solution: 


C 


Exercise: 


Problem: Where is the inferior oblique muscle located? 


a. in the abdomen 

b. in the eye socket 

c. in the anterior neck 
d. in the face 


Solution: 
B 
Exercise: 
Problem: What is the action of the masseter? 


a. swallowing 

b. chewing 

c. moving the lips 
d. closing the eye 


Solution: 

B 
Exercise: 

Problem:The names of the extrinsic tongue muscles commonly end in 
a. -glottis 
b. -glossus 
c. -gluteus 
d. -hyoid 

Solution: 

B 
Exercise: 


Problem: What is the function of the erector spinae? 


a. movement of the arms 

b. stabilization of the pelvic girdle 
c. postural support 

d. rotating of the vertebral column 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem:Explain the difference between axial and appendicular muscles. 


Solution: 


Axial muscles originate on the axial skeleton (the bones in the head, neck, and core of the body), 
whereas appendicular muscles originate on the bones that make up the body’s limbs. 


Exercise: 


Problem: Describe the muscles of the anterior neck. 
Solution: 


The muscles of the anterior neck are arranged to facilitate swallowing and speech. They work on the 
hyoid bone, with the suprahyoid muscles pulling up and the infrahyoid muscles pulling down. 


Exercise: 


Problem: Why are the muscles of the face different from typical skeletal muscle? 
Solution: 


Most skeletal muscles create movement by actions on the skeleton. Facial muscles are different in 
that they create facial movements and expressions by pulling on the skin—no bone movements are 
involved. 


Glossary 


anterior scalene 
a muscle anterior to the middle scalene 


appendicular 
of the arms and legs 


axial 
of the trunk and head 


buccinator 
muscle that compresses the cheek 


corrugator supercilii 
prime mover of the eyebrows 


deglutition 
swallowing 


digastric 
muscle that has anterior and posterior bellies and elevates the hyoid bone and larynx when one 
swallows; it also depresses the mandible 


epicranial aponeurosis 
(also, galea aponeurosis) flat broad tendon that connects the frontalis and occipitalis 


erector spinae group 
large muscle mass of the back; primary extensor of the vertebral column 


extrinsic eye muscles 
originate outside the eye and insert onto the outer surface of the white of the eye, and create eyeball 
movement 


frontalis 
front part of the occipitofrontalis muscle 


genioglossus 
muscle that originates on the mandible and allows the tongue to move downward and forward 


geniohyoid 
muscle that depresses the mandible, and raises and pulls the hyoid bone anteriorly 


hyoglossus 
muscle that originates on the hyoid bone to move the tongue downward and flatten it 


iliocostalis cervicis 
muscle of the iliocostalis group associated with the cervical region 


iliocostalis group 
laterally placed muscles of the erector spinae 


iliocostalis lumborum 
muscle of the iliocostalis group associated with the lumbar region 


iliocostalis thoracis 
muscle of the iliocostalis group associated with the thoracic region 


infrahyoid muscles 
anterior neck muscles that are attached to, and inferior to the hyoid bone 


lateral pterygoid 
muscle that moves the mandible from side to side 


longissimus capitis 
muscle of the longissimus group associated with the head region 


longissimus cervicis 
muscle of the longissimus group associated with the cervical region 


longissimus group 
intermediately placed muscles of the erector spinae 


longissimus thoracis 
muscle of the longissimus group associated with the thoracic region 


masseter 
main muscle for chewing that elevates the mandible to close the mouth 


mastication 
chewing 


medial pterygoid 
muscle that moves the mandible from side to side 


middle scalene 
longest scalene muscle, located between the anterior and posterior scalenes 


multifidus 
muscle of the lumbar region that helps extend and laterally flex the vertebral column 


mylohyoid 
muscle that lifts the hyoid bone and helps press the tongue to the top of the mouth 


occipitalis 
posterior part of the occipitofrontalis muscle 


occipitofrontalis 
muscle that makes up the scalp with a frontal belly and an occipital belly 


omohyoid 
muscle that has superior and inferior bellies and depresses the hyoid bone 


orbicularis oculi 
circular muscle that closes the eye 


orbicularis oris 
circular muscle that moves the lips 


palatoglossus 
muscle that originates on the soft palate to elevate the back of the tongue 


posterior scalene 
smallest scalene muscle, located posterior to the middle scalene 


scalene muscles 
flex, laterally flex, and rotate the head; contribute to deep inhalation 


segmental muscle group 
interspinales and intertransversarii muscles that bring together the spinous and transverse processes of 
each consecutive vertebra 


semispinalis capitis 
transversospinales muscle associated with the head region 


semispinalis cervicis 
transversospinales muscle associated with the cervical region 


semispinalis thoracis 
transversospinales muscle associated with the thoracic region 


spinalis capitis 
muscle of the spinalis group associated with the head region 


spinalis cervicis 
muscle of the spinalis group associated with the cervical region 


spinalis group 
medially placed muscles of the erector spinae 


spinalis thoracis 
muscle of the spinalis group associated with the thoracic region 


splenius 
posterior neck muscles; includes the splenius capitis and splenius cervicis 


splenius capitis 
neck muscle that inserts into the head region 


splenius cervicis 
neck muscle that inserts into the cervical region 


sternocleidomastoid 
major muscle that laterally flexes and rotates the head 


sternohyoid 
muscle that depresses the hyoid bone 


sternothyroid 
muscle that depresses the larynx’s thyroid cartilage 


styloglossus 
muscle that originates on the styloid bone, and allows upward and backward motion of the tongue 


stylohyoid 
muscle that elevates the hyoid bone posteriorly 


suprahyoid muscles 
neck muscles that are superior to the hyoid bone 


temporalis 
muscle that retracts the mandible 


thyrohyoid 
muscle that depresses the hyoid bone and elevates the larynx’s thyroid cartilage 


transversospinales 
muscles that originate at the transverse processes and insert at the spinous processes of the vertebrae 


Muscles of the Abdominal Wall and Thorax 
By the end of this section, you will be able to: 


e Identify the intrinsic skeletal muscles of the back and neck, and the skeletal muscles of the abdominal 
wall and thorax 

e Identify the movement and function of the intrinsic skeletal muscles of the back and neck, and the 
skeletal muscles of the abdominal wall and thorax 


It is a complex job to balance the body on two feet and walk upright. The muscles of the vertebral column, 
thorax, and abdominal wall extend, flex, and stabilize different parts of the body’s trunk. The deep muscles of 
the core of the body help maintain posture as well as carry out other functions. The brain sends out electrical 
impulses to these various muscle groups to control posture by alternate contraction and relaxation. This is 
necessary so that no single muscle group becomes fatigued too quickly. If any one group fails to function, 
body posture will be compromised. 


Muscles of the Abdomen 


There are four pairs of abdominal muscles that cover the anterior and lateral abdominal region and meet at the 
anterior midline. These muscles of the anterolateral abdominal wall can be divided into four groups: the 
external obliques, the internal obliques, the transversus abdominis, and the rectus abdominis ([link] and 
(link). 

Muscles of the Abdomen 


Pectoralis major External oblique 


Latissimus dorsi Rectus Transversus 


sheath abdominis 
Anterior serratus muscles 


External oblique 


Linea alba (of the 
rectus sheath) 


Rectus abdominis 
(enclosed within 
rectus sheath) 


Tendinous intersections 
(between the anterior \ ea 

segments of the rectus } Rectus abdominis ; 
abdominis) Aponeurosis of 
internal oblique 


Internal oblique 


Quadratus lumborum 


llia of hip bones 


lliacus 


Sacrum 


Psoas major 


youy 


(a) The anterior abdominal muscles include the medially 


located rectus abdominis, which is covered by a sheet of 
connective tissue called the rectus sheath. On the flanks of 

the body, medial to the rectus abdominis, the abdominal 

wall is composed of three layers. The external oblique 
muscles form the superficial layer, while the internal 

oblique muscles form the middle layer, and the transverses 
abdominus forms the deepest layer. (b) The muscles of the 
lower back move the lumbar spine but also assist in femur 


movements. 
Muscles of the Abdomen 
Target 
motion Prime 
Movement Target direction mover Origin Insertion 
Satan External Ribs Ribs 7— 
Twisting at waist; also Vertebral P . obliques; ; 10; linea 
: ; lateral ; 5-12; 
bending to the side column ; internal Ba alba; 
flexion ° ilium a 
obliques ilium 
Squeezing abdomen nea Sternum; 
; F Tlium; : 
during forceful Abdominal ; Transversus . linea 
: ; : Compression ; ribs 5— 
exhalations, defecation, cavity abdominus 10 alba; 
urination, and childbirth pubis 
Sternum; 
Ea ] é R F f ; 
Sitting up vee Flexion oe ; Pubis ribs 5 
column abdominis 
and 7 
. . Vertebral Lateral Quadratus lium; Ri 
Bending to the side : ribs 5— vertebrae 
column flexion lumborum 10 LLL4 


There are three flat skeletal muscles in the antero-lateral wall of the abdomen. The external oblique, closest to 
the surface, extend inferiorly and medially, in the direction of sliding one’s four fingers into pants pockets. 
Perpendicular to it is the intermediate internal oblique, extending superiorly and medially, the direction the 
thumbs usually go when the other fingers are in the pants pocket. The deep muscle, the transversus 
abdominis, is arranged transversely around the abdomen, similar to the front of a belt on a pair of pants. This 
arrangement of three bands of muscles in different orientations allows various movements and rotations of the 
trunk. The three layers of muscle also help to protect the internal abdominal organs in an area where there is 
no bone. 


The linea alba is a white, fibrous band that is made of the bilateral rectus sheaths that join at the anterior 
midline of the body. These enclose the rectus abdominis muscles (a pair of long, linear muscles, commonly 
called the “sit-up” muscles) that originate at the pubic crest and symphysis, and extend the length of the body’s 
trunk. Each muscle is segmented by three transverse bands of collagen fibers called the tendinous 


intersections. This results in the look of “six-pack abs,” as each segment hypertrophies on individuals at the 
gym who do many sit-ups. 


The posterior abdominal wall is formed by the lumbar vertebrae, parts of the ilia of the hip bones, psoas major 
and iliacus muscles, and quadratus lumborum muscle. This part of the core plays a key role in stabilizing the 
rest of the body and maintaining posture. 


Note: 

Career Connections 

Physical Therapists 

Those who have a muscle or joint injury will most likely be sent to a physical therapist (PT) after seeing their 
regular doctor. PTs have a master’s degree or doctorate, and are highly trained experts in the mechanics of 
body movements. Many PTs also specialize in sports injuries. 

If you injured your shoulder while you were kayaking, the first thing a physical therapist would do during 
your first visit is to assess the functionality of the joint. The range of motion of a particular joint refers to the 
normal movements the joint performs. The PT will ask you to abduct and adduct, circumduct, and flex and 
extend the arm. The PT will note the shoulder’s degree of function, and based on the assessment of the injury, 
will create an appropriate physical therapy plan. 

The first step in physical therapy will probably be applying a heat pack to the injured site, which acts much 
like a warm-up to draw blood to the area, to enhance healing. You will be instructed to do a series of exercises 
to continue the therapy at home, followed by icing, to decrease inflammation and swelling, which will 
continue for several weeks. When physical therapy is complete, the PT will do an exit exam and send a 
detailed report on the improved range of motion and return of normal limb function to your doctor. Gradually, 
as the injury heals, the shoulder will begin to function correctly. A PT works closely with patients to help 
them get back to their normal level of physical activity. 


Muscles of the Thorax 
The muscles of the chest serve to facilitate breathing by changing the size of the thoracic cavity ([link]). When 


you inhale, your chest rises because the cavity expands. Alternately, when you exhale, your chest falls because 
the thoracic cavity decreases in size. 


Muscles of the Thorax 


Target motion Prime 
Movement Target direction mover Origin Insertion 
Sternum; 
Inhalation; Thoracic Compression; . ribs 6-12; Central 
: : ; Diaphragm 
exhalation cavity expansion lumbar tendon 


vertebrae 


Muscles of the Thorax 


Target motion Prime 
Movement Target direction mover Origin Insertion 
Rib Rib 
Elevation superior inferior to 
F : d External 
Inhalation;exhalation Ribs (expands : to each each 
: : intercostals é ; 
thoracic cavity) intercostal intercostal 
muscle muscle 
Movement Rib Rib 
along inferi : 
superior/inferior Internal cna cia eaieg 
Forced exhalation Ribs : : : each to each 
axis to bring intercostals : : 
. intercostal intercostal 
ribs closer 
muscle muscle 
together 
The Diaphragm 


The change in volume of the thoracic cavity during breathing is due to the alternate contraction and relaxation 
of the diaphragm ((link]). It separates the thoracic and abdominal cavities, and is dome-shaped at rest. The 
superior surface of the diaphragm is convex, creating the elevated floor of the thoracic cavity. The inferior 
surface is concave, creating the curved roof of the abdominal cavity. 

Muscles of the Diaphragm 


Central tendon 
of diaphragm - —— Sternum 


Vena cava 
passing through 
caval opening 


Esophagus 


passing through 
esophageal hiatus 


Aorta passing 
through aortic 
hiatus 


12th (floating) ribs 


Left psoas major 


Left quadratus 
lumborum 


Vertebrae 


Diaphragm (inferior view) 


The diaphragm separates the thoracic and 
abdominal cavities. 


Defecating, urination, and even childbirth involve cooperation between the diaphragm and abdominal muscles 
(this cooperation is referred to as the “Valsalva maneuver”). You hold your breath by a steady contraction of 
the diaphragm; this stabilizes the volume and pressure of the peritoneal cavity. When the abdominal muscles 
contract, the pressure cannot push the diaphragm up, so it increases pressure on the intestinal tract 
(defecation), urinary tract (urination), or reproductive tract (childbirth). 


The inferior surface of the pericardial sac and the inferior surfaces of the pleural membranes (parietal pleura) 
fuse onto the central tendon of the diaphragm. To the sides of the tendon are the skeletal muscle portions of the 
diaphragm, which insert into the tendon while having a number of origins including the xiphoid process of the 
sternum anteriorly, the inferior six ribs and their cartilages laterally, and the lumbar vertebrae and 12th ribs 
posteriorly. 


The diaphragm also includes three openings for the passage of structures between the thorax and the abdomen. 
The inferior vena cava passes through the caval opening, and the esophagus and attached nerves pass through 
the esophageal hiatus. The aorta, thoracic duct, and azygous vein pass through the aortic hiatus of the posterior 
diaphragm. 


The Intercostal Muscles 


There are three sets of muscles, called intercostal muscles, which span each of the intercostal spaces. The 
principal role of the intercostal muscles is to assist in breathing by changing the dimensions of the rib cage 
(link). 

Intercostal Muscles 


Clavicle Gi - af 


Ribs = 


Pectoralis minor 


Innermost 
intercostal 


Pectoralis major 
(dissected) 


Internal 
intercostal 


Sternum 


Serratus 
anterior 


External —_ oe 
intercostals — 


alt 


intercostal 


Internal 
intercostal 


The external intercostals are located laterally on the sides of the body. 
The internal intercostals are located medially near the sternum. The 
innermost intercostals are located deep to both the internal and external 
intercostals. 


The 11 pairs of superficial external intercostal muscles aid in inspiration of air during breathing because 
when they contract, they raise the rib cage, which expands it. The 11 pairs of internal intercostal muscles, 
just under the externals, are used for expiration because they draw the ribs together to constrict the rib cage. 
The innermost intercostal muscles are the deepest, and they act as synergists for the action of the internal 
intercostals. 


Muscles of the Pelvic Floor and Perineum 


The pelvic floor is a muscular sheet that defines the inferior portion of the pelvic cavity. The pelvic 
diaphragm, spanning anteriorly to posteriorly from the pubis to the coccyx, comprises the levator ani and the 
ischiococcygeus. Its openings include the anal canal and urethra, and the vagina in women. 


The large levator ani consists of two skeletal muscles, the pubococcygeus and the iliococcygeus ((link]). The 
levator ani is considered the most important muscle of the pelvic floor because it supports the pelvic viscera. It 
resists the pressure produced by contraction of the abdominal muscles so that the pressure is applied to the 
colon to aid in defecation and to the uterus to aid in childbirth (assisted by the ischiococcygeus, which pulls 
the coccyx anteriorly). This muscle also creates skeletal muscle sphincters at the urethra and anus. 

Muscles of the Pelvic Floor 


Pubic crest 


Vaginal canal 
(females only) 


Rectal canal 


Sacrum 


lliac crests 


Pelvic diaphragm (superior view) 


The pelvic floor muscles support the pelvic 
organs, resist intra-abdominal pressure, and work 
as sphincters for the urethra, rectum, and vagina. 


The perineum is the diamond-shaped space between the pubic symphysis (anteriorly), the coccyx 
(posteriorly), and the ischial tuberosities (laterally), lying just inferior to the pelvic diaphragm (levator ani and 
coccygeus). Divided transversely into triangles, the anterior is the urogenital triangle, which includes the 
external genitals. The posterior is the anal triangle, which contains the anus ([link]). The perineum is also 
divided into superficial and deep layers with some of the muscles common to men and women ([(link]). 
Women also have the compressor urethrae and the sphincter urethrovaginalis, which function to close the 
vagina. In men, there is the deep transverse perineal muscle that plays a role in ejaculation. 

Muscles of the Perineum 


Penis 


Clitoris 


Ischiocavernosus 


Bulbospongiosus (aka 
bulbocavernosus) 


Urethra 


Vagina 


Transverse perineal 
muscles 


Anus 


External anal sphincter 
Levator ani 
Coccyx 


Gluteus maximus 


Male perineal muscles: inferior view Female perineal muscles: inferior view 


The perineum muscles play roles in urination in both sexes, 
ejaculation in men, and vaginal contraction in women. 


Muscles of the Perineum Common to Men and Women 


Target motion ; 
[tarot [Taman | Primemover | ova | insertion | 


Movement 


Defecation; Abdominal Superior Levator ani Pubis; ischium Urethra; anal 
urination; birth; cavity (resists pubococcygeus; canal; perineal 
coughing pressure levator ani body; coccyx 


during iliococcygeus 
abdominal 
compression) 


Superficial muscles 


None— Perineal body Superficial Ischium Perineal body 


supports transverse 
perineal body perineal 
maintaining 

anus at center 


of perineum 


Involuntary Urethra Compression Bulbospongiosus | Perineal body Perineal 
response that membrane; 
compresses corpus 

urethra when spongiosum 
excreting urine of penis; deep 
in both sexes or fascia of penis; 
while ejaculating clitoris in 

in males; also female 

aids in erection of 
penis in males 


Compresses Veins of penis Compression Ischiocavernosus | Ischium; ischial | Pubic 

veins to maintain | and clitoris rami; pubic rami | symphysis; 
erection of penis corpus 

in males; erection cavernosum of 
of clitoris in penis in male; 
females clitoris of 


female 


Deep muscles 


Voluntarily Urethra Compression External urethral Ischial rami; Male: median 
compresses sphincter pubic rami raphe; female: 
urethra during vaginal wall 

urination 


External anal 
sphincter 


Closes anus Sphincter Anoccoccygeal | Perineal body 


ligament 


Chapter Review 


Made of skin, fascia, and four pairs of muscle, the anterior abdominal wall protects the organs located in the 
abdomen and moves the vertebral column. These muscles include the rectus abdominis, which extends through 
the entire length of the trunk, the external oblique, the internal oblique, and the transversus abdominus. The 


quadratus lumborum forms the posterior abdominal wall. 


The muscles of the thorax play a large role in breathing, especially the dome-shaped diaphragm. When it 
contracts and flattens, the volume inside the pleural cavities increases, which decreases the pressure within 
them. As a result, air will flow into the lungs. The external and internal intercostal muscles span the space 
between the ribs and help change the shape of the rib cage and the volume-pressure ratio inside the pleural 


cavities during inspiration and expiration. 


The perineum muscles play roles in urination in both sexes, ejaculation in men, and vaginal contraction in 
women. The pelvic floor muscles support the pelvic organs, resist intra-abdominal pressure, and work as 


sphincters for the urethra, rectum, and vagina. 


Review Questions 


Exercise: 


Problem: Which of the following abdominal muscles is not a part of the anterior abdominal wall? 


a. quadratus lumborum 
b. rectus abdominis 

c. interior oblique 

d. exterior oblique 


Solution: 
A 
Exercise: 
Problem: Which muscle pair plays a role in respiration? 


a. intertransversarii, interspinales 

b. semispinalis cervicis, semispinalis thoracis 
c. trapezius, rhomboids 

d. diaphragm, scalene 


Solution: 
D 
Exercise: 
Problem: What is the linea alba? 


a. a small muscle that helps with compression of the abdominal organs 
b. a long tendon that runs down the middle of the rectus abdominis 

c. a long band of collagen fibers that connects the hip to the knee 

d. another name for the tendinous inscription 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


Describe the fascicle arrangement in the muscles of the abdominal wall. How do they relate to each 
other? 


Solution: 


Arranged into layers, the muscles of the abdominal wall are the internal and external obliques, which run 
on diagonals, the rectus abdominis, which runs straight down the midline of the body, and the transversus 
abdominis, which wraps across the trunk of the body. 


Exercise: 


Problem: What are some similarities and differences between the diaphragm and the pelvic diaphragm? 


Solution: 


Both diaphragms are thin sheets of skeletal muscle that horizontally span areas of the trunk. The 
diaphragm separating the thoracic and abdominal cavities is the primary muscle of breathing. The pelvic 


diaphragm, consisting of two paired muscles, the coccygeus and the levator ani, forms the pelvic floor at 
the inferior end of the trunk. 


Glossary 


anal triangle 
posterior triangle of the perineum that includes the anus 


caval opening 
opening in the diaphragm that allows the inferior vena cava to pass through; foramen for the vena cava 


compressor urethrae 
deep perineal muscle in women 


deep transverse perineal 
deep perineal muscle in men 


diaphragm 
skeletal muscle that separates the thoracic and abdominal cavities and is dome-shaped at rest 


external intercostal 
superficial intercostal muscles that raise the rib cage 


external oblique 
superficial abdominal muscle with fascicles that extend inferiorly and medially 


iliococcygeus 
muscle that makes up the levator ani along with the pubococcygeus 


innermost intercostal 
the deepest intercostal muscles that draw the ribs together 


intercostal muscles 
muscles that span the spaces between the ribs 


internal intercostal 
muscles the intermediate intercostal muscles that draw the ribs together 


internal oblique 
flat, intermediate abdominal muscle with fascicles that run perpendicular to those of the external oblique 


ischiococcygeus 
muscle that assists the levator ani and pulls the coccyx anteriorly 


levator ani 
pelvic muscle that resists intra-abdominal pressure and supports the pelvic viscera 


linea alba 
white, fibrous band that runs along the midline of the trunk 


pelvic diaphragm 
muscular sheet that comprises the levator ani and the ischiococcygeus 


perineum 
diamond-shaped region between the pubic symphysis, coccyx, and ischial tuberosities 


pubococcygeus 
muscle that makes up the levator ani along with the iliococcygeus 


quadratus lumborum 
posterior part of the abdominal wall that helps with posture and stabilization of the body 


rectus abdominis 
long, linear muscle that extends along the middle of the trunk 


rectus sheaths 
tissue that makes up the linea alba 


sphincter urethrovaginalis 
deep perineal muscle in women 


tendinous intersections 
three transverse bands of collagen fibers that divide the rectus abdominis into segments 


transversus abdominis 
deep layer of the abdomen that has fascicles arranged transversely around the abdomen 


urogenital triangle 
anterior triangle of the perineum that includes the external genitals 


Muscles of the Pectoral Girdle and Upper Limbs 
By the end of this section, you will be able to: 


e Identify the muscles of the pectoral girdle and upper limbs 
e Identify the movement and function of the pectoral girdle and upper limbs 


Muscles of the shoulder and upper limb can be divided into four groups: muscles that stabilize and position the 
pectoral girdle, muscles that move the arm, muscles that move the forearm, and muscles that move the wrists, 
hands, and fingers. The pectoral girdle, or shoulder girdle, consists of the lateral ends of the clavicle and scapula, 
along with the proximal end of the humerus, and the muscles covering these three bones to stabilize the shoulder 
joint. The girdle creates a base from which the head of the humerus, in its ball-and-socket joint with the glenoid 
fossa of the scapula, can move the arm in multiple directions. 


Muscles That Position the Pectoral Girdle 


Muscles that position the pectoral girdle are located either on the anterior thorax or on the posterior thorax ([link] 
and [link]). The anterior muscles include the subclavius, pectoralis minor, and serratus anterior. The posterior 
muscles include the trapezius, rhomboid major, and rhomboid minor. When the rhomboids are contracted, your 
scapula moves medially, which can pull the shoulder and upper limb posteriorly. 

Muscles That Position the Pectoral Girdle 


Deltoid (cut) 
Coracoid process 

of scapula 

Pectoralis major 

(cut) Acromion process Rhomboid 
Scapula of scapula minor 


Subclavius 


Pectoralis minor Rhomboid 


major 
Sternum Serratus : 


anterior 


Deltoid 


Trapezius 


Pectoral girdle muscle (left anterior lateral view) Pectoral girdle muscles (posterior view) 


The muscles that stabilize the pectoral girdle make it a steady 

base on which other muscles can move the arm. Note that the 

pectoralis major and deltoid, which move the humerus, are cut 
here to show the deeper positioning muscles. 


Muscles that Position the Pectoral Girdle 


Position Target 
in the motion Prime 
thorax Movement Target direction mover Origin Insertion 
Stabilizes ; 
Anterior clavicle durin ia 
6 Clavicle Depression Subclavius First rib surface of 
thorax movement by Ears 


depressing it 


Muscles that Position the Pectoral Girdle 


Position 
in the 
thorax 


Anterior 
thorax 


Anterior 
thorax 


Posterior 
thorax 


Posterior 
thorax 


Posterior 
thorax 


Movement 


Rotates 
shoulder 
anteriorly 
(throwing 
motion); assists 
with inhalation 


Moves arm 
from side of 
body to front 
of body; assists 
with inhalation 


Elevates 
shoulders 
(shrugging); 
pulls shoulder 
blades 
together; tilts 
head 
backwards 


Stabilizes 
scapula during 
pectoral girdle 
movement 


Stabilizes 
scapula during 
pectoral girdle 
movement 


Target 


Scapula; 
ribs 


Scapula; 
ribs 


Scapula; 
cervical 
spine 


Scapula 


Scapula 


Muscles That Move the Humerus 


Similar to the muscles that position the pectoral girdle, muscles that cross the shoulder joint and move the humerus 


Target 
motion 
direction 


Scapula: 
depresses; 
ribs: elevates 


Scapula: 
protracts; 
ribs: elevates 


Scapula: 
rotests 
inferiorly, 
retracts, 
elevates, and 
depresses; 
spine: 
extends 


Retracts; 
rotates 
inferiorly 


Retracts; 
rotates 
inferiorly 


Prime 
mover 


Pectoralis 
minor 


Serratus 
anterior 


Trapezius 


Rhomboid 
major 


Rhomboid 
minor 


Origin 


Anterior 
surfaces 
of 
certain 
ribs (2-4 
or 3-5) 


Muscle 
slips 
from 
certain 
ribs (1-8 
or 1-9) 


Skull; 
vertebral 
column 


Thoracic 
vertebrae 
(T2-T5) 


Cervical 
and 
thoracic 
vertebrae 
(C7 and 
T1) 


Insertion 


Coracoid 
process of 
scapula 


Anterior 
surface of 
vertebral 
border of 
scapula 


Acromion 
and spine 
of 
scapula; 
clavicle 


Medial 
border of 
scapula 


Medial 
border of 
scapula 


bone of the arm include both axial and scapular muscles ((link] and [link]). The two axial muscles are the 
pectoralis major and the latissimus dorsi. The pectoralis major is thick and fan-shaped, covering much of the 
superior portion of the anterior thorax. The broad, triangular latissimus dorsi is located on the inferior part of the 
back, where it inserts into a thick connective tissue shealth called an aponeurosis. 


Muscles That Move the Humerus 


Pectoralis 
major 


Latissimus dorsi 


AM 


(a) Pectoralis major and latissimus dorsi 
(left anterior lateral view) 


(b) Left deltoid and left latissimus dorsi 
(posterior view) 


Teres minor Supraspinatus 


Deltoid 


(cut) Spine of 


NN scapula 
| 
Deltoid (cut) 


Coracoid process 
of scapula 
Infraspinatus 


Pectoralis Humerus 


major (cut) 
Subscapularis 
Teres major 
Teres major Latissimus dorsi 

(near its origin) 
Serratus Triceps brachii: long head 
anterior 


Triceps brachii: lateral head 


B Ya 


(d) Deep muscles of the left shoulder 
(posterior view) 


(c) Deep muscles of the left shoulder 
(anterior lateral view) 


(a, c) The muscles that move the humerus anteriorly are 
generally located on the anterior side of the body and 
originate from the sternum (e.g., pectoralis major) or the 
anterior side of the scapula (e.g., subscapularis). (b) The 
muscles that move the humerus superiorly generally 
originate from the superior surfaces of the scapula and/or 
the clavicle (e.g., deltoids). The muscles that move the 
humerus inferiorly generally originate from middle or 
lower back (e.g., latissiumus dorsi). (d) The muscles that 
move the humerus posteriorly are generally located on the 
posterior side of the body and insert into the scapula (e.g., 
infraspinatus). 


Muscles That Move the Humerus 


Target motion 
direction 


Movement Target Prime mover Origin Insertion 


Axial muscles 


Brings elbows Humerus Flexion; Pectoralis Clavicle; sternum; | Greater 
together; moves adduction; major cartilage of certain | tubercle of 
elbow up (as medial ribs (1-6 or 1-7); | humerus 
during an uppercut rotation aponeurosis of 
punch) external oblique 

muscle 
Moves elbow back Humerus; Humerus: Latissimus Thoracic Intertubercular 
(as in elbowing scapula extension, dorsi vertebrae sulcus of 
someone standing adduction, and (T7-T12); lumbar | humerus 
behind you); medial rotation; vertebrae; lower 
spreads elbows scapula: ribs (9-12); 


apart depression iliac crest 


Scapular muscles 


Lifts arms at Humerus Abduction; Deltoid Trapezius; Deltoid 
shoulder flexion; clavicle; tuberosity 
extension; acromion; of humerus 
medial and spine of scapula 
lateral rotation 


Assists pectoralis 
major in bringing 
elbows together 


Humerus Medial Subscapularis Subscapular Lesser 
rotation fossa of tubercle of 
scapula humerus 
shoulder joint during 
movement of the 
pectoral girdle 


and stabilizes 

Rotates elbow Humerus Abduction Supraspinatus Supraspinous Greater 
outwards, as during fossa of scapula | tubercle of 
a tennis swing humerus 
Rotates elbow Humerus Extension; Infraspinatus Infraspinous fossa | Greater 
outwards, as during adduction of scapula tubercle of 
a tennis swing humerus 
Assists infraspinatus | Humerus Extension; Teres major Posterior surface | Intertubercular 
in rotating elbow adduction of scapula sulcus of 
outwards humerus 
Assists infraspinatus | Humerus Extension; Teres minor Lateral border of Greater 

in rotating elbow adduction dorsal scapular tubercle of 
outwards surface humerus 


Moves elbow up Humerus Flexion; Coracobra Coracoid process | Medial surface 
and across body, adduction chialis of scapula of humerus, 

as when putting shaft 

hand on chest 


The rest of the shoulder muscles originate on the scapula. The anatomical and ligamental structure of the shoulder 
joint and the arrangements of the muscles covering it, allows the arm to carry out different types of movements. 
The deltoid, the thick muscle that creates the rounded lines of the shoulder is the major abductor of the arm, but it 
also facilitates flexing and medial rotation, as well as extension and lateral rotation. The subscapularis originates 
on the anterior scapula and medially rotates the arm. Named for their locations, the supraspinatus (superior to the 
spine of the scapula) and the infraspinatus (inferior to the spine of the scapula) abduct the arm, and laterally rotate 
the arm, respectively. The thick and flat teres major is inferior to the teres minor and extends the arm, and assists 
in adduction and medial rotation of it. The long teres minor laterally rotates and extends the arm. Finally, the 
coracobrachialis flexes and adducts the arm. 


The tendons of the deep subscapularis, supraspinatus, infraspinatus, and teres minor connect the scapula to the 
humerus, forming the rotator cuff (musculotendinous cuff), the circle of tendons around the shoulder joint. When 
baseball pitchers undergo shoulder surgery it is usually on the rotator cuff, which becomes pinched and inflamed, 
and may tear away from the bone due to the repetitive motion of bring the arm overhead to throw a fast pitch. 


Muscles That Move the Forearm 


The forearm, made of the radius and ulna bones, has four main types of action at the hinge of the elbow joint: 
flexion, extension, pronation, and supination. The forearm flexors include the biceps brachii, brachialis, and 
brachioradialis. The extensors are the triceps brachii and anconeus. The pronators are the pronator teres and the 
pronator quadratus, and the supinator is the only one that turns the forearm anteriorly. When the forearm faces 
anteriorly, it is supinated. When the forearm faces posteriorly, it is pronated. 


The biceps brachii, brachialis, and brachioradialis flex the forearm. The two-headed biceps brachii crosses the 
shoulder and elbow joints to flex the forearm, also taking part in supinating the forearm at the radioulnar joints and 
flexing the arm at the shoulder joint. Deep to the biceps brachii, the brachialis provides additional power in 
flexing the forearm. Finally, the brachioradialis can flex the forearm quickly or help lift a load slowly. These 
muscles and their associated blood vessels and nerves form the anterior compartment of the arm (anterior flexor 
compartment of the arm) ([link] and [link]). 


Muscles That Move the Forearm 


Biceps brachii 
(short head) Triceps brachii 


. ¥ (lateral head) 
Biceps brachii 


(long head) 


Triceps brachii 
(long head) 


Left upper arm muscles (anterior lateral view) Left upper arm muscles (posterior view) 


Triceps brachii 
Brachioradialis 


Extensor carpi radialis longus 


Extensor carpi radialis brevis 
Lateral 

epicondyle 
of humerus 


Pronator teres 
Abductor pollicis longus 
Extensor pollicis brevis 

Extensor pollicis 


Flexor carpi 
radialis 

. Anconeus 
Palmaris longus 


longus Extensor carpi 
Flexor carpi ulnaris ulnaris 
Extensor digitorum 
Flexor digitorum superficialis Extensor digiti 
i minimi 
g 
Left forearm superficial muscles (palmar view) Left forearm superficial muscles (dorsal view) 


Lateral epicondyle of humerus 
Medial epicondyle of humerus 
Supinator 


Flexor pollicis longus 


Brachialis Abductor pollicis longus . 
(cut) \ Medial 
epicondyle 


Pronator quadratus of humerus 


Flexor digitorum Extensor pollicis longus 


profundus Fl 
4 4 jexor 
Extensor pollicis Be digitorum 


brevis é : profundus 


Flexor carpi ulnaris 


Flexor retinaculum 
(cut) 


Extensor indicis 


y Extensor 
rz - retinaculum 


Left forearm deep muscles (palmar view) Left forearm deep muscles (dorsal view) 


The muscles originating in the upper arm flex, extend, pronate, and 
supinate the forearm. The muscles originating in the forearm move the 
wrists, hands, and fingers. 


Muscles That Move the Forearm 


Target motion ‘ 
aie | taroet | Tagetcnane | Primemover | ovgin | insertion | 


Anterior muscles (flexion) 


Performs a bicep Forearm Flexion; Biceps brachii Coracoid process; | Radial 

curl; also allows supination tubercle above tuberosity 

palm of hand to glenoid cavity 

point toward body 

while flexing 

Forearm Flexion Brachialis Front of distal Coronoid 

humerus process of ulna 

Assists and Forearm Flexion Brachioradialis Lateral Base of styloid 

stabilizes elbow supracondylar process of 

during bicep-curl tidge at distal end | radius 

motion of humerus 


Posterior muscles (extension) 


Extends forearm, Forearm Extension Triceps brachii Infraglenoid Olecranon 
as during a punch tubercle of process of ulna 
scapula; posterior 
shaft of humerus; 
posterior humeral 
shaft distal to 
radial groove 


Assists in extending | Forearm Extension; Anconeus Lateral epicondyle | Lateral aspect 
forearm; also allows abduction of humerus of olecranon 
forearm to extend process of ulna 
away from body 


Anterior muscles (pronation) 


Turns hand Forearm Pronation Pronator teres 
palm-down 

Assists in Forearm Pronation Pronator Distal portion Distal surface 
turning hand quadratus of anterior ulnar of anterior 
palm-down shaft radius 


Posterior muscles (supination) 


Medial epicondyle | Lateral radius 
of humerus; 

coronoid process 

of ulna 


Turns hand Forearm Supination Supinator Lateral epicondyle | Proximal end 
palm-up of humerus; of radius 
proximal ulna 


Muscles That Move the Wrist, Hand, and Fingers 


Wrist, hand, and finger movements are facilitated by two groups of muscles. The forearm is the origin of the 
extrinsic muscles of the hand. The palm is the origin of the intrinsic muscles of the hand. 


Muscles of the Arm That Move the Wrists, Hands, and Fingers 


The muscles in the anterior compartment of the forearm (anterior flexor compartment of the forearm) originate 
on the humerus and insert onto different parts of the hand. These make up the bulk of the forearm. From lateral to 
medial, the superficial anterior compartment of the forearm includes the flexor carpi radialis, palmaris 
longus, flexor carpi ulnaris, and flexor digitorum superficialis. The flexor digitorum superficialis flexes the 
hand as well as the digits at the knuckles, which allows for rapid finger movements, as in typing or playing a 
musical instrument (see [link] and [link]). However, poor ergonomics can irritate the tendons of these muscles as 
they slide back and forth with the carpal tunnel of the anterior wrist and pinch the median nerve, which also travels 
through the tunnel, causing Carpal Tunnel Syndrome. The deep anterior compartment produces flexion and 
bends fingers to make a fist. These are the flexor pollicis longus and the flexor digitorum profundus. 


The muscles in the superficial posterior compartment of the forearm (superficial posterior extensor 
compartment of the forearm) originate on the humerus. These are the extensor radialis longus, extensor carpi 
radialis brevis, extensor digitorum, extensor digiti minimi, and the extensor carpi ulnaris. 


The muscles of the deep posterior compartment of the forearm (deep posterior extensor compartment of the 
forearm) originate on the radius and ulna. These include the abductor pollicis longus, extensor pollicis brevis, 
extensor pollicis longus, and extensor indicis (see [link]). 

Muscles That Move the Wrist, Hands, and Forearm 


Target motion 


Target direction 


Superficial anterior compartment of forearm 


Eine mene | ori 


Insertion 


Bends wrist toward body; tilts | Wrist; 
hand to side away from body | hand 


Flexion; 
abduction 


Assists in bending hand up Flexion 


toward shoulder 


Flexor carpi 
radialis 


Medial epicondyle 
of humerus 


Palmaris 
longus 


Medial epicondyle 
of humerus 


Base of second 
and third 
metacarpals 


Palmar 
aponeurosis; skin 
and fascia of palm 


Assists in bending hand up 
toward shoulder; tilts hand to 
side away from body; 
stabilizes wrist 


Flexion, 
abduction 


also bends wrist toward body | fingers 


Bends fingers to make a fist | Wrist; Flexion 
fingers 

Deep anterior compartment of forearm 

Bends tip of Thumb Flexion 

thumb 

Bends fingers to make a fist; | Wrist; Flexion 


Superficial posterior compartment of forearm 


Wrist Extension; 


abduction 


Straightens wrist away from 
body; tilts hand to side away 
from body 


Assists extensor radialis 


Extension, 


longus in extending and abduction 


Flexor carpi 
ulnaris 


Medial epicondyle 

of humerus; olecranon 
process; posterior 
surface of ulna 


Flexor digitorum 
superficialis 


Medial epicondyle of 
humerus; coronoid 
process of ulna; shaft 
of radius 


Anterior surface of 
radius; interosseous 
membrane 


Flexor pollicis 
longus 


Flexor digitorum | Coronoid process; 


profundus anteromedial surface of 
ulna; interosseous 
membrane 

Extensor Lateral supracondylar 


radialis longus ridge of humerus 


Extensor carpi 
radialis brevis 


Lateral epicondyle 
of humerus 


abducting wrist; also 
stabilizes hand during finger 


flexion. 


Opens fingers and moves Wrist; Extension; 

them sideways away from fingers abduction 

the body 

Extends little finger Little Extension 
finger 

Straightens wrist away from | Wrist Extension; 

body; tilts hand to side adduction 


toward body 
Deep posterior compartment of forearm 


Thumb: 
abduction, 


Moves thumb sideways 
toward body; extends thumb; 
moves hand sideways 
toward body 


extension; 
wrist: abduction 


Extends thumb Thumb Extension 
Extends thumb Thumb Extension 
Extends index finger; Wrist; Extension 
straightens wrist away from index 
body finger 


Extensor Lateral epicondyle 
digitorum of humerus 
Extensor Lateral epicondyle 
digiti minimi of humerus 


Extensor carpi 
ulnaris 


Lateral epicondyle of 
humerus; posterior 
border of ulna 


Abductor Posterior surface 

pollicis longus of radius and ulna; 
interosseous 
membrane 


Extensor Dorsal shaft of radius 

pollicis brevis and ulna; interosseous 
membrane 

Extensor Dorsal shaft of radius 

pollicis longus and ulna; interosseous 


membrane 


Posterior surface 
of distal ulna; 
interosseous membrane 


Extensor indicis 


Pisiform, hamate 
bones, and base of 
fifth metacarpal 


Middle phalanges 
of fingers 2-5 


Distal phalanx 
of thumb 


Distal phalanges 
of fingers 2-5 


Base of second 
metacarpal 


Base of third 
metacarpal 


Extensor 
expansions; 
distal phalanges 
of fingers 


Extensor 
expansion; 
distal phalanx of 
finger 5 


Base of fifth 
metacarpal 


Base of first 
metacarpal; 
trapezium 


Base of proximal 
phalanx of thumb 


Base of distal 
phalanx of thumb 


Tendon of extensor 
digitorum of index 
finger 


The tendons of the forearm muscles attach to the wrist and extend into the hand. Fibrous bands called retinacula 
sheath the tendons at the wrist. The flexor retinaculum extends over the palmar surface of the hand while the 


extensor retinaculum extends over the dorsal surface of the hand. 


Intrinsic Muscles of the Hand 


The intrinsic muscles of the hand both originate and insert within it ([link]). These muscles allow your fingers to 
also make precise movements for actions, such as typing or writing. These muscles are divided into three groups. 
The thenar muscles are on the radial aspect of the palm. The hypothenar muscles are on the medial aspect of the 


palm, and the intermediate muscles are midpalmar. 


The thenar muscles include the abductor pollicis brevis, opponens pollicis, flexor pollicis brevis, and the 
adductor pollicis. These muscles form the thenar eminence, the rounded contour of the base of the thumb, and 
all act on the thumb. The movements of the thumb play an integral role in most precise movements of the hand. 


The hypothenar muscles include the abductor digiti minimi, flexor digiti minimi brevis, and the opponens 
digiti minimi. These muscles form the hypothenar eminence, the rounded contour of the little finger, and as 
such, they all act on the little finger. Finally, the intermediate muscles act on all the fingers and include the 


lumbrical, the palmar interossei, and the dorsal interossei. 


Intrinsic Muscles of the Hand 


Vee 


Opponens pollicis 


Abductor digiti 
minimi Abductor pollicis 
brevl 

Flexor digiti ie 
minimi brevis 


Pisometacarpal 


Flexor pollicis ligament 
brevis 


FLX ‘I : Opponens digiti 
; 2 minimi 
! Adductor pollicis oe 


f 
y 
{ | 
Lumbricalis 
muscles U 


Superficial muscles of left hand (palmar) 


Dorsal interossei 
muscles 


Palmar interossei 
muscles 


Interossei muscles of left hand (palmar view) Interossei muscles of left hand (dorsal view) 
The intrinsic muscles of the hand both originate and 
insert within the hand. These muscles provide the fine 
motor control of the fingers by flexing, extending, 
abducting, and adducting the more distal finger and 
thumb segments. 


Intrinsic Muscles of the Hand 


Target 
motion Prime 
Muscle Movement Target direction mover 
Thenar Moves thumb . eeu 
Thumb Abduction pollicis 
muscles toward body . 
brevis 
Moves thumb 
Thenar across palm to ve Opponens 
Th ice 
muscles touch other nae Pepesiien pollicis 


fingers 


Origin 


Flexor 
retinaculum; 
and nearby 
carpals 


Flexor 
retinaculum; 
trapezium 


Insertion 


Lateral 
base of 
proximal 
phalanx o 
thumb 


Anterior ¢ 
first 
metacarpe 


Intrinsic Muscles of the Hand 


Muscle 
Thenar 


muscles 


Thenar 
muscles 


Hypothenar 
muscles 


Hypothenar 
muscles 


Hypothenar 
muscles 


Intermediate 
muscles 


Movement 


Flexes thumb 


Moves thumb 
away from 
body 


Moves little 
finger toward 
body 


Flexes little 
finger 


Moves little 
finger across 
palm to touch 
thumb 


Flexes each 
finger at 
metacarpo- 
phalangeal 


joints; extends 


each finger at 


interphalangeal 


joints 


Target 


Thumb 


Thumb 


Little 
finger 


Little 
finger 


Little 
finger 


Fingers 


Target 
motion 
direction 


Flexion 


Adduction 


Abduction 


Flexion 


Opposition 


Flexion 


Prime 
mover 


Flexor 
pollicis 
brevis 


Adductor 
pollicis 


Abductor 
digiti 
minimi 


Flexor 
digiti 
minimi 
brevis 


Opponens 
digiti 
minimi 


Lumbricals 


Origin 


Flexor 
retinaculum; 
trapezium 


Capitate 
bone; bases 
of 
metacarpals 
2-4; front of 
metacarpal 

3 


Pisiform 
bone 


Hamate 
bone; flexor 
retinaculum 


Hamate 
bone; flexor 
retinaculum 


Palm 
(lateral sides 
of tendons 
in flexor 
digitorum 
profundus) 


Insertion 


Lateral 
base of 
proximal 
phalanx o 
thumb 


Medial 
base of 
proximal 
phalanx o 
thumb 


Medial 
side of 
proximal 
phalanx o 
little finge 


Medial 
side of 
proximal 
phalanx o 
little finge 


Medial 
side of 
fifth 
metacarpé 


Fingers 2- 
5 (lateral 
edges of 
extension: 
expansion 
on first 
phalanges 


Intrinsic Muscles of the Hand 


Target 
motion Prime 
Muscle Movement Target direction mover Origin Insertion 
Adducts and : Peters! 
Side of each expansion 
flexes each : 
; metacarpal on first 
Bngenat that faces hal 
oe phalanx o 
‘ metacarpo- Adduction; f 
Intermediate : an Palmar metacarpal each finge 
phalangeal Fingers flexion; . ; 
muscles eeeeen : interossei 3 (absent (except 
joints; extends extension ; 
é from finger 3) 
each finger at : 
; metacarpal on side 
interphalangeal 3) facing 
join : 
JoMts finger 3 
Abducts and ee ae 
flexes the three foi eee i 
middle fingers Sian 
at metacarpo- finger 
5 halangeal Ab ion; : : 
Intermediate pe nge ‘ dnctic Dorsal Sides of extensor 
joints; extends Fingers flexion; . . : 
muscles : interossei metacarpals expansion 
the three extension sen tict 
middle fingers phalanx o 
a ; 
a side 
interphalangeal opposite 
join 3 
jeans finger 3 


Chapter Review 


The clavicle and scapula make up the pectoral girdle, which provides a stable origin for the muscles that move the 
humerus. The muscles that position and stabilize the pectoral girdle are located on the thorax. The anterior thoracic 
muscles are the subclavius, pectoralis minor, and the serratus anterior. The posterior thoracic muscles are the 
trapezius, levator scapulae, rhomboid major, and rhomboid minor. Nine muscles cross the shoulder joint to move 
the humerus. The ones that originate on the axial skeleton are the pectoralis major and the latissimus dorsi. The 
deltoid, subscapularis, supraspinatus, infraspinatus, teres major, teres minor, and coracobrachialis originate on the 
scapula. 


The forearm flexors include the biceps brachii, brachialis, and brachioradialis. The extensors are the triceps brachii 
and anconeus. The pronators are the pronator teres and the pronator quadratus. The supinator is the only one that 
turns the forearm anteriorly. 


The extrinsic muscles of the hands originate along the forearm and insert into the hand in order to facilitate crude 
movements of the wrists, hands, and fingers. The superficial anterior compartment of the forearm produces 
flexion. These muscles are the flexor carpi radialis, palmaris longus, flexor carpi ulnaris, and the flexor digitorum 
superficialis. The deep anterior compartment produces flexion as well. These are the flexor pollicis longus and the 
flexor digitorum profundus. The rest of the compartments produce extension. The extensor carpi radialis longus, 
extensor carpi radialis brevis, extensor digitorum, extensor digiti minimi, and extensor carpi ulnaris are the 
muscles found in the superficial posterior compartment. The deep posterior compartment includes the abductor 
longus, extensor pollicis brevis, extensor pollicis longus, and the extensor indicis. 


Finally, the intrinsic muscles of the hands allow our fingers to make precise movements, such as typing and 
writing. They both originate and insert within the hand. The thenar muscles, which are located on the lateral part of 


the palm, are the abductor pollicis brevis, opponens pollicis, flexor pollicis brevis, and adductor pollicis. The 
hypothenar muscles, which are located on the medial part of the palm, are the abductor digiti minimi, flexor digiti 
minimi brevis, and opponens digiti minimi. The intermediate muscles, located in the middle of the palm, are the 
lumbricals, palmar interossei, and dorsal interossei. 


Review Questions 


Exercise: 


Problem:The rhomboid major and minor muscles are deep to the 


a. rectus abdominis 

b. scalene muscles 

c. trapezius 

d. ligamentum nuchae 


Solution: 


C 


Exercise: 


Problem:Which muscle extends the forearm? 


a. biceps brachii 
b. triceps brachii 
c. brachialis 

d. deltoid 


Solution: 


B 


Exercise: 


Problem: What is the origin of the wrist flexors? 


a. the lateral epicondyle of the humerus 
b. the medial epicondyle of the humerus 
c. the carpal bones of the wrist 

d. the deltoid tuberosity of the humerus 


Solution: 


B 


Exercise: 


Problem:Which muscles stabilize the pectoral girdle? 


a. axial and scapular 

b. axial 

c. appendicular 

d. axial and appendicular 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem:The tendons of which muscles form the rotator cuff? Why is the rotator cuff important? 


Solution: 


Tendons of the infraspinatus, supraspinatus, teres minor, and the subscapularis form the rotator cuff, which 
forms a foundation on which the arms and shoulders can be stabilized and move. 


Exercise: 


Problem: List the general muscle groups of the shoulders and upper limbs as well as their subgroups. 


Solution: 


The muscles that make up the shoulders and upper limbs include the muscles that position the pelvic girdle, 
the muscles that move the humerus, the muscles that move the forearm, and the muscles that move the wrists, 
hands, and fingers. 


Glossary 


abductor digiti minimi 
muscle that abducts the little finger 


adductor pollicis 
muscle that adducts the thumb 


abductor pollicis brevis 
muscle that abducts the thumb 


abductor pollicis longus 
muscle that inserts into the first metacarpal 


anconeus 
small muscle on the lateral posterior elbow that extends the forearm 


anterior compartment of the arm 
(anterior flexor compartment of the arm) the biceps brachii, brachialis, brachioradialis, and their associated 
blood vessels and nerves 


anterior compartment of the forearm 
(anterior flexor compartment of the forearm) deep and superficial muscles that originate on the humerus and 
insert into the hand 


biceps brachii 
two-headed muscle that crosses the shoulder and elbow joints to flex the forearm while assisting in supinating 
it and flexing the arm at the shoulder 


brachialis 


muscle deep to the biceps brachii that provides power in flexing the forearm. 


brachioradialis 
muscle that can flex the forearm quickly or help lift a load slowly 


coracobrachialis 
muscle that flexes and adducts the arm 


deep anterior compartment 
flexor pollicis longus, flexor digitorum profundus, and their associated blood vessels and nerves 


deep posterior compartment of the forearm 
(deep posterior extensor compartment of the forearm) the abductor pollicis longus, extensor pollicis brevis, 
extensor pollicis longus, extensor indicis, and their associated blood vessels and nerves 


deltoid 
shoulder muscle that abducts the arm as well as flexes and medially rotates it, and extends and laterally 
rotates it 


dorsal interossei 
muscles that abduct and flex the three middle fingers at the metacarpophalangeal joints and extend them at the 
interphalangeal joints 


extensor carpi radialis brevis 
muscle that extends and abducts the hand at the wrist 


extensor carpi ulnaris 
muscle that extends and adducts the hand 


extensor digiti minimi 
muscle that extends the little finger 


extensor digitorum 
muscle that extends the hand at the wrist and the phalanges 


extensor indicis 
muscle that inserts onto the tendon of the extensor digitorum of the index finger 


extensor pollicis brevis 
muscle that inserts onto the base of the proximal phalanx of the thumb 


extensor pollicis longus 
muscle that inserts onto the base of the distal phalanx of the thumb 


extensor radialis longus 
muscle that extends and abducts the hand at the wrist 


extensor retinaculum 
band of connective tissue that extends over the dorsal surface of the hand 


extrinsic muscles of the hand 
muscles that move the wrists, hands, and fingers and originate on the arm 


flexor carpi radialis 
muscle that flexes and abducts the hand at the wrist 


flexor carpi ulnaris 
muscle that flexes and adducts the hand at the wrist 


flexor digiti minimi brevis 
muscle that flexes the little finger 


flexor digitorum profundus 
muscle that flexes the phalanges of the fingers and the hand at the wrist 


flexor digitorum superficialis 
muscle that flexes the hand and the digits 


flexor pollicis brevis 
muscle that flexes the thumb 


flexor pollicis longus 
muscle that flexes the distal phalanx of the thumb 


flexor retinaculum 
band of connective tissue that extends over the palmar surface of the hand 


hypothenar 
group of muscles on the medial aspect of the palm 


hypothenar eminence 
rounded contour of muscle at the base of the little finger 


infraspinatus 
muscle that laterally rotates the arm 


intermediate 
group of midpalmar muscles 


intrinsic muscles of the hand 
muscles that move the wrists, hands, and fingers and originate in the palm 


latissimus dorsi 
broad, triangular axial muscle located on the inferior part of the back 


lumbrical 
muscle that flexes each finger at the metacarpophalangeal joints and extend each finger at the interphalangeal 
joints 

opponens digiti minimi 


muscle that brings the little finger across the palm to meet the thumb 


opponens pollicis 
muscle that moves the thumb across the palm to meet another finger 


palmar interossei 
muscles that abduct and flex each finger at the metacarpophalangeal joints and extend each finger at the 
interphalangeal joints 


palmaris longus 
muscle that provides weak flexion of the hand at the wrist 


pectoral girdle 
shoulder girdle, made up of the clavicle and scapula 


pectoralis major 
thick, fan-shaped axial muscle that covers much of the superior thorax 


pectoralis minor 
muscle that moves the scapula and assists in inhalation 


pronator quadratus 
pronator that originates on the ulna and inserts on the radius 


pronator teres 
pronator that originates on the humerus and inserts on the radius 


retinacula 
fibrous bands that sheath the tendons at the wrist 


rhomboid major 
muscle that attaches the vertebral border of the scapula to the spinous process of the thoracic vertebrae 


rhomboid minor 
muscle that attaches the vertebral border of the scapula to the spinous process of the thoracic vertebrae 


rotator cuff 
(also, musculotendinous cuff) the circle of tendons around the shoulder joint 


serratus anterior 
large and flat muscle that originates on the ribs and inserts onto the scapula 


subclavius 
muscle that stabilizes the clavicle during movement 


subscapularis 
muscle that originates on the anterior scapula and medially rotates the arm 


superficial anterior compartment of the forearm 
flexor carpi radialis, palmaris longus, flexor carpi ulnaris, flexor digitorum superficialis, and their associated 
blood vessels and nerves 


superficial posterior compartment of the forearm 
extensor radialis longus, extensor carpi radialis brevis, extensor digitorum, extensor digiti minimi, extensor 
carpi ulnaris, and their associated blood vessels and nerves 


supinator 
muscle that moves the palm and forearm anteriorly 


supraspinatus 
muscle that abducts the arm 


teres major 
muscle that extends the arm and assists in adduction and medial rotation of it 


teres minor 
muscle that laterally rotates and extends the arm 


thenar 
group of muscles on the lateral aspect of the palm 


thenar eminence 
rounded contour of muscle at the base of the thumb 


trapezius 
muscle that stabilizes the upper part of the back 


triceps brachii 
three-headed muscle that extends the forearm 


Muscles of the Pelvic Girdle and Lower Limbs 
By the end of this section, you will be able to: 


e Identify the appendicular muscles of the pelvic girdle and lower limb 
e Identify the movement and function of the pelvic girdle and lower limb 


The appendicular muscles of the lower body position and stabilize the 
pelvic girdle, which serves as a foundation for the lower limbs. 
Comparatively, there is much more movement at the pectoral girdle than at 
the pelvic girdle. There is very little movement of the pelvic girdle because 
of its connection with the sacrum at the base of the axial skeleton. The 
pelvic girdle is less range of motion because it was designed to stabilize and 
support the body. 


Muscles of the Thigh 


What would happen if the pelvic girdle, which attaches the lower limbs to 
the torso, were capable of the same range of motion as the pectoral girdle? 
For one thing, walking would expend more energy if the heads of the 
femurs were not secured in the acetabula of the pelvis. The body’s center of 
gravity is in the area of the pelvis. If the center of gravity were not to 
remain fixed, standing up would be difficult as well. Therefore, what the leg 
muscles lack in range of motion and versatility, they make up for in size and 
power, facilitating the body’s stabilization, posture, and movement. 


Gluteal Region Muscles That Move the Femur 


Most muscles that insert on the femur (the thigh bone) and move it, 
originate on the pelvic girdle. The psoas major and iliacus make up the 
iliopsoas group. Some of the largest and most powerful muscles in the 
body are the gluteal muscles or gluteal group. The gluteus maximus is the 
largest; deep to the gluteus maximus is the gluteus medius, and deep to the 
gluteus medius is the gluteus minimus, the smallest of the trio ((link] and 
[link]). 

Hip and Thigh Muscles 


Quadratus 
lumborum 


Psoas major 


lliacus ; 
Pectineus 


ihn Sacrum 

Tensor Adductor longus 

fascia latae Gracilis 
Adductor 

Rectus magnus 

femoris Sartorius 

Vastus Vastus medialis 


lateralis eer tendon 
(or patellar tendon) 
i Patellar ligament 


Superficial pelvic and Bee muscles 
of right leg (anterior view) 


Crest of ilium Gluteus 
medius (cut) 
i Sacrum 
lliac ; Gluteus 
crest Pectineus roe minimus 
: urator 

o : Pubis interns Piriformis 

Cbturator Glluteus Superior 

externus rasanlis gemellus 


Inferior 
gemellus 


Obturator 
externus 


Adductor (cut) 
brevis 

Adductor Adductor 
longus group 
Adductor Gracilis 
magnus 


Quadratus 
‘ femoris 

Semimembranosus 
Biceps 


Semitendinosus femoris 


Deep pelvic and thigh muscles Pelvic and thigh muscles of 
of right leg (anterior view) right leg (posterior view) 


The large and powerful muscles of the hip that 
move the femur generally originate on the pelvic 
girdle and insert into the femur. The muscles that 

move the lower leg typically originate on the 
femur and insert into the bones of the knee joint. 
The anterior muscles of the femur extend the lower 
leg but also aid in flexing the thigh. The posterior 
muscles of the femur flex the lower leg but also 
aid in extending the thigh. A combination of 
gluteal and thigh muscles also adduct, abduct, and 
rotate the thigh and lower leg. 


Gluteal Region Muscles That Move the Femur 


Target motion A 7 
| __ Movement | Tarot | T™gescgne” | Primemover | origin | serton | 


lliopsoas group 


Raises knee at hip, as if 
performing a knee attack; 
assists lateral rotators in 
twisting thigh (and lower leg) 
outward; assists with bending 
over, maintaining posture 


Raises knee at hip, as if 
performing a knee attack; 
assists lateral rotators in 
twisting thigh (and lower leg) 
outward; assists with bending 
over, maintaining posture 


Gluteal group 


Lowers knee and moves 
thigh back, as when getting 
ready to kick a ball 


Opens thighs, as when 
doing a split 


Brings the thighs back 
together 


Assists with raising knee at 
hip and opening thighs; 
maintains posture by 
stabilizing the iliotibial track, 
which connects to the knee 


Lateral rotators 


Twists thigh (and lower leg) 
outward; maintains posture 
by stabilizing hip joint 


Twists thigh (and lower leg) 
outward; maintains posture 
by stabilizing hip joint 


Twists thigh (and lower leg) 
outward; maintains posture 
by stabilizing hip joint 


Twists thigh (and lower leg) 
outward; maintains posture 
by stabilizing hip joint 


Twists thigh (and lower 
leg) outward; maintains 
posture by stabilizing 

hip joint 

Twists thigh (and lower leg) 
outward; maintains posture 
by stabilizing hip joint 


Adductors 


Brings the thighs back 
together; assists with 
raising the knee 


Brings the thighs back 
together; assists with 
raising the knee 


Brings the thighs back 
together; assists with raising 
the knee and moving the 
thigh back 


Opens thighs; assists with 
raising the knee and turning 
the thigh (and lower leg) 
inward 


Lumbar vertebrae 
(L1-L5); 
thoracic vertebra (T12) 


Psoas major 


lliacus 


Gluteus maximus 


Thigh: flexion and 
lateral rotation; 
torso: flexion 


lliac fossa; iliac crest; 
lateral sacrum 


Thigh: flexion and 
lateral rotation; 
torso: flexion 


Extension 


Dorsal ilium; sacrum; 
coccyx 


Femur 


Femur 


Gluteus 


minimus 


Femur 


Flexion; 
abduction 


Tensor fascia 
lata 


Obutrator 
internus 


Femur Anterior aspect of 
iliac crest; anterior 


superior iliac spine 


Anterolateral surface 
of sacrum 


Lateral rotation 


Inner surface of 
obturator membrane; 
greater sciatic notch; 
margins of obturator 
foramen 


Lateral rotation 


Outer surfaces of 
obturator membrane, 
pubic, and ischium; 
margins of obturator 
foramen 


Obturator 
externus 


Lateral rotation 


Lateral rotation Superior 


gemellus 


Ischial spine 


Inferior 
gemellus 


Lateral rotation Ischial tuberosity 


Quadratus 
femoris 


Lateral rotation Ischial tuberosity 


Adduction; 
flexion 


Adductor longus | Pubis near pubic 


symphysis 


Adduction; Adductor brevis 


flexion 


Body of pubis; inferior 
ramus of pubis 


Adductor 
magnus 


Adduction; 
flexion; 
extension 


Ischial rami; pubic rami; 
ischial tuberosity 


Adduction; flexion; | Pectineus Pectineal line of pubis 


medial rotation 


Lesser trochanter 
of femur 


Lesser trochanter 
of femur 


Gluteal tuberosity 
of femur; iliotibial 
tract 


Abduction Gluteus Lateral surface of illum | Greater trochanter 
medius of femur 

External surface of Greater trochanter 

ilium of femur 


lliotibial tract 


Greater trochanter 
of femur 


Greater trochanter 
in front of piriformis 


Trochanteric fossa 
of posterior femur 


Greater trochanter 
of femur 


Greater trochanter 
of femur 


Trochanteric 
crest of femur 


Linea aspera 


Linea aspera 
above adductor 
longus 


Linea aspera; 
adductor tubercle 
of femur 


Lesser trochanter 
to linea aspera of 
posterior aspect of 
femur 


The tensor fascia latae is a thick, squarish muscle in the superior aspect of 
the lateral thigh. It acts as a synergist of the gluteus medius and iliopsoas in 
flexing and abducting the thigh. It also helps stabilize the lateral aspect of 
the knee by pulling on the iliotibial tract (band), making it taut. Deep to 
the gluteus maximus, the piriformis, obturator internus, obturator 
externus, superior gemellus, inferior gemellus, and quadratus femoris 
laterally rotate the femur at the hip. 


The adductor longus, adductor brevis, and adductor magnus can both 
medially and laterally rotate the thigh depending on the placement of the 
foot. The adductor longus flexes the thigh, whereas the adductor magnus 
extends it. The pectineus adducts and flexes the femur at the hip as well. 
The pectineus is located in the femoral triangle, which is formed at the 

junction between the hip and the leg and also includes the femoral nerve, 
the femoral artery, the femoral vein, and the deep inguinal lymph nodes. 


Thigh Muscles That Move the Femur, Tibia, and Fibula 


Deep fascia in the thigh separates it into medial, anterior, and posterior 
compartments (see [link] and [link]). The muscles in the medial 
compartment of the thigh are responsible for adducting the femur at the 
hip. Along with the adductor longus, adductor brevis, adductor magnus, and 
pectineus, the strap-like gracilis adducts the thigh in addition to flexing the 
leg at the knee. 

Thigh Muscles That Move the Femur, Tibia, and Fibula 


Target Target metion Prime mover 
direction 


Medial compartment of thigh 


Moves back of lower 
legs up toward 
buttocks, as when 
kneeling; assists in 
opening thighs 


Femur; 
tibia/fibula 


Tibia/fibula: 
flexion; thigh: 
adduction 


Gracilis 


Anterior compartment of thigh: Quadriceps femoris group 


Moves lower leg out 
in front of body, as 
when kicking; assists 
in raising the knee 


Moves lower leg out 
in front of body, as 
when kicking 


Moves lower leg out 
in front of body, as 
when kicking 


Moves lower leg out 
in front of body, as 
when kicking 


Moves back of lower 
legs up and back 
toward the buttocks, 
as when kneeling; 
assists in moving 
thigh diagonally 
upward and outward 
as when mounting 

a bike 


Posterior compartment of thigh: Hamstring group 


Moves back of lower 
legs up and back 
oward the buttocks, 
as when kneeling; 
moves thigh down 
and back; twists the 
high (and lower leg) 
outward 


Moves back of lower 
legs up toward 
buttocks, as when 
kneeling; moves 

high down and back; 
twists the thigh 

(and lower leg) 
inward 


Moves back of lower 
legs up and back 
toward the buttocks 
as when kneeling; 
moves thigh down 
and back; twists the 
thigh (and lower leg) 
inward 


Femur; 
tibia/fibula 


Tibia/fibula 


Tibia/fibula 


Tibia/fibula 


Femur; 
tibia/fibula 


Femur; 
tibia/fibula 


Femur; 
tibia/fibula 


Femur; 
tibia/fibula 


Tibia/fibula: 
extension; 
thigh: flexion 


Extension 


Extension 


Extension 


Tibia: flexion; 
thigh: flexion, 
abduction, 
lateral 
rotation 


Tibia/fibula: 
flexion; thigh: 
extension, 
lateral 
rotation 


Tibia/fibula: 
flexion; thigh: 
extension, 
medial 
rotation 


Tibia/fibula: 
flexion; thigh: 
extension, 
medial 
rotation 


Rectus 
femoris 


Vastus 
lateralis 


Vastus 
medialis 


Vastus 
intermedius 


Sartorius 


Biceps 
femoris 


Semitendinosus 


Semi- 
membranosus 


Inferior ramus; 
body of pubis; 
ischial ramus 


Anterior inferior 
iliac spine; superior 
margin of 
acetabulum 


Greater trochanter; 
intertrochanteric 
line; linea aspera 


Linea aspera; 
intertrochanteric 
line 


Proximal femur 
shaft 


Anterior superior 
iliac spine 


Ischial tuberosity; 
linea aspera; 
distal femur 


Ischial tuberosity 


Ischial tuberosity 


Medial surface 
of tibia 


Patella; tibial 
tuberosity 


Patella; tibial 
tuberosity 


Patella; tibial 
tuberosity 


Patella; tibial 
tuberosity 


Medial aspect 
of proximal 
tibia 


Head of fibula; 
lateral condyle 
of tibia 


Upper tibial 
shaft 


Medial condyle 
of tibia; lateral 
condyle of 
femur 


The muscles of the anterior compartment of the thigh flex the thigh and 
extend the leg. This compartment contains the quadriceps femoris group, 
which actually comprises four muscles that extend and stabilize the knee. 
The rectus femoris is on the anterior aspect of the thigh, the vastus 
lateralis is on the lateral aspect of the thigh, the vastus medialis is on the 


medial aspect of the thigh, and the vastus intermedius is between the 
vastus lateralis and vastus medialis and deep to the rectus femoris. The 
tendon common to all four is the quadriceps tendon (patellar tendon), 
which inserts into the patella and continues below it as the patellar 
ligament. The patellar ligament attaches to the tibial tuberosity. In addition 
to the quadriceps femoris, the sartorius is a band-like muscle that extends 
from the anterior superior iliac spine to the medial side of the proximal 
tibia. This versatile muscle flexes the leg at the knee and flexes, abducts, 
and laterally rotates the leg at the hip. This muscle allows us to sit cross- 
legged. 


The posterior compartment of the thigh includes muscles that flex the leg 
and extend the thigh. The three long muscles on the back of the knee are the 
hamstring group, which flexes the knee. These are the biceps femoris, 
semitendinosus, and semimembranosus. The tendons of these muscles 
form the popliteal fossa, the diamond-shaped space at the back of the knee. 


Muscles That Move the Feet and Toes 


Similar to the thigh muscles, the muscles of the leg are divided by deep 
fascia into compartments, although the leg has three: anterior, lateral, and 
posterior ([link] and [link]). 

Muscles of the i Leg 


Gastrocnemius 
(lateral head) 


Superior extensor" 
retinaculum 


Inferior edeneare—— 
retinaculum 


Superficial muscles of the right 
lower leg (anterior view) 


Tibialis anterior 
Fibularis longus 
Extensor digitorum 
longus 

Fibularis brevis 
Extensor hallucis 
longus 

Fibularis tertius 


Gastrocnemius 
(medial head) 


Plantaris 


Soleus 


Calcaneal (Achilles) | 
tendon } | 
> 


Superficial muscles of the right 
lower leg (posterior view) 


Calcaneus (heel) 


Popliteus 


Soleus (cut) 
Fibularis longus 
Tibialis posterior 


Flexor digitorum 
longus 


Flexor hallucis 
longus 


Fibularis brevis 


Deep muscles of the right 
lower leg (posterior view) 


The muscles of the anterior compartment of the lower leg are 
generally responsible for dorsiflexion, and the muscles of the 
posterior compartment of the lower leg are generally 
responsible for plantar flexion. The lateral and medial 
muscles in both compartments invert, evert, and rotate the 
foot. 


Muscles That Move the Feet and Toes 


Target motion ‘ 
Movement | Target | Taegeamenon | Primemover | origin] serton | 


Anterior compartment of leg 


Raises the sole of the foot off the 
ground, as when preparing to 
foot-tap; bends the inside of the 
foot upwards, as when catching 
your balance while falling laterally 
toward the opposite side as the 
balancing foot 


Raises the sole of the foot off 
the ground, as when preparing 
to foot-tap; extends the big toe 


Raises the sole of the foot off 
the ground, as when preparing 
to foot-tap; extends toes 


Lateral compartment of leg 


Lowers the sole of the foot to 

the ground, as when foot-tapping 
or jumping; bends the inside of 
the foot downwards, as when 
catching your balance while falling 
laterally toward the same side 

as the balancing foot 


Lowers the sole of the foot to 

the ground, as when foot-tapping 
or jumping; bends the inside of 
the foot downward, as when 
catching your balance while 
falling laterally toward the same 
side as the balancing foot 


Foot; 
big toe 


Foot; 
toes 
2-5 


Dorsiflexion; 
inversion 


Foot: dorsiflexion; 
big toe: extension 


Foot: dorsiflexion; 
toes: extension 


Plantar flexion 
and eversion 


Plantar flexion 
and eversion 


Posterior compartment of leg: Superficial muscles 


Lowers the sole of the foot to the 
ground, as when foot-tapping or 
jumping; assists in moving the 
back of the lower legs up and 
back toward the buttocks 


Lowers the sole of the foot to the 
ground, as when foot-tapping or 
jumping; maintains posture while 
walking 


Lowers the sole of the foot to the 
ground, as when foot-tapping or 
jumping; assists in moving the 


back of the lower legs up and back 


toward the buttocks 


Lowers the sole of the foot to the 
ground, as when foot-tapping or 
jumping 


Posterior compartment of leg: Deep muscles 


Moves the back of the lower legs 
up and back toward the buttocks; 
assists in rotation of the leg at the 
knee and thigh 


Lowers the sole of the foot to the 
ground, as when foot-tapping or 
jumping; bends the inside of the 
foot upward and flexes toes 


Flexes the big toe 


Foot; 
tibia/ 
fibula 


Foot; 
tibia/ 
fibula 


Tibia/ 
fibula 
Foot; 
toes 2-5 
Big toe; 
foot 


Foot: plantar 
flexion; 
tibia/fibula: flexion 


Plantar 
flexion 


Foot: plantar 
flexion; 
tibia/fibula: 
flexion 


Plantar flexion 


Tibialis anterior 


Extensor hallucis 
longus 


Extensor 
digitorum longus 


Fibularis longus 


Fibularis 
(peroneus) brevis 


Gastrocnemius 


Plantaris 


Tibialis posterior 


Tibia/fibula: flexion} Popliteus 


thigh and lower 
leg: medial and 
lateral rotation 


Foot: plantar 
flexion and 
inversion 
toes: flexion 


Big toe: 
flexion foot: 
plantar flexion 


Flexor digitorum 
longus 


Flexor hallucis 
longus 


Lateral condyle 
and upper tibial 
shaft; interosseous 
membrane 


Anteromedial fibula 
shaft; interosseous 
membrane 


Lateral condyle of 
tibia; proximal portion 
of fibula; interosseous 
membrane 


Upper portion of 
lateral fibula 


Distal fibula shaft 


Medial and lateral 
condyles of femur 


Superior tibia; 
fibula; interosseous 
membrane 


Posterior femur 
above lateral 
condyle 


Superior tibia and 
fibula; interosseous 
membrane 


Lateral condyle of 
femur; lateral 
meniscus 


Posterior tibia 


Midshaft of fibula; 
interosseous 
membrane 


Interior surface 
of medial 
cuneiform; 

First metatarsal 
bone 


Distal phalanx 
of big toe 


Middle and distal 
phalanges of 
toes 2-5 


First metatarsal; 
medial cuneiform 


Proximal end 
of fifth 
metatarsal 


Posterior 
calcaneus 


Posterior 
calcaneus 


Calcaneus or 
calcaneus 
tendon 


Several tarsals 
and metatarsals 
2-4 


Proximal tibia 


Distal phalanges 
of toes 2-5 


Distal phalanx 
of big toe 


The muscles in the anterior compartment of the leg: the tibialis anterior, 
a long and thick muscle on the lateral surface of the tibia, the extensor 
hallucis longus, deep under it, and the extensor digitorum longus, lateral 


to it, all contribute to raising the front of the foot when they contract. The 
fibularis tertius, a small muscle that originates on the anterior surface of 
the fibula, is associated with the extensor digitorum longus and sometimes 
fused to it, but is not present in all people. Thick bands of connective tissue 
called the superior extensor retinaculum (transverse ligament of the 
ankle) and the inferior extensor retinaculum, hold the tendons of these 
muscles in place during dorsiflexion. 


The lateral compartment of the leg includes two muscles: the fibularis 
longus (peroneus longus) and the fibularis brevis (peroneus brevis). The 
superficial muscles in the posterior compartment of the leg all insert onto 
the calcaneal tendon (Achilles tendon), a strong tendon that inserts into the 
calcaneal bone of the ankle. The muscles in this compartment are large and 
strong and keep humans upright. The most superficial and visible muscle of 
the calf is the gastrocnemius. Deep to the gastrocnemius is the wide, flat 
soleus. The plantaris runs obliquely between the two; some people may 
have two of these muscles, whereas no plantaris is observed in about seven 
percent of other cadaver dissections. The plantaris tendon is a desirable 
substitute for the fascia lata in hernia repair, tendon transplants, and repair 
of ligaments. There are four deep muscles in the posterior compartment of 
the leg as well: the popliteus, flexor digitorum longus, flexor hallucis 
longus, and tibialis posterior. 


The foot also has intrinsic muscles, which originate and insert within it 
(similar to the intrinsic muscles of the hand). These muscles primarily 
provide support for the foot and its arch, and contribute to movements of 
the toes ([link] and [link]). The principal support for the longitudinal arch of 
the foot is a deep fascia called plantar aponeurosis, which runs from the 
calcaneus bone to the toes (inflammation of this tissue is the cause of 
“plantar fasciitis,” which can affect runners. The intrinsic muscles of the 
foot consist of two groups. The dorsal group includes only one muscle, the 
extensor digitorum brevis. The second group is the plantar group, which 
consists of four layers, starting with the most superficial. 

Intrinsic Muscles of the Foot 


Tendocalcaneus 


Fibularis longus Extensor digitorum brevis 


——_____ Tibialis anterior 
Extensor digitorum longus 
-. Extensor hallucis longus 
Y 


Fibularis brevis 


Fibularis tertius 


(a) Dorsal superficial muscles of 
the right foot (lateral view) 


Abductor 
digiti 
minimi 


Plantar 
aponeurosis 


Quadratus Flexor digiti 
a plantae minimi brevis 
allucis 
Flexor Flexor 
digitorum hallucis 
brevis ae brevis 
y @ ti) i (Vy . ; 
Cl} Wh \\\/ \ Wi Lumbricals 
v dH W 
(b) Superficial muscles of the (c) Intermediate muscles of (d) Deep muscles of the 
left sole (plantar view) the left sole (plantar view) left sole (plantar view) 


The muscles along the dorsal side of the foot (a) generally 
extend the toes while the muscles of the plantar side of the 
foot (b, c, d) generally flex the toes. The plantar muscles 
exist in three layers, providing the foot the strength to 
counterbalance the weight of the body. In this diagram, 
these three layers are shown from a plantar view beginning 
with the bottom-most layer just under the plantar skin of 
the foot (b) and ending with the top-most layer (d) located 
just inferior to the foot and toe bones. 


Intrinsic Muscles in the Foot 


Target motion Prime 


Dorsal group 


Extends toes 2-5 Toes 2-5 | Extension Extensor Calcaneus; Base of proximal 
digitorum extensor phalanx of big toe; 
brevis retinaculum extensor expansions 

on toes 2-5 


Plantar group (layer 1) 


Abducts and flexes Adduction; Abductor Calcaneal Proximal phalanx of 
big toe flexion hallucis tuberosity; flexor big toe 
retinaculum 


Flexes toes 2-4 Middle Flexion Flexor Calcaneal Middle phalanx of 
toes digitorum tuberosity toes 2-4 
brevis 
Abducts and flexes Toe 5 Abduction; Abductor Calcaneal tuberosity | Proximal phalanx 
small toe flexion digiti minimi of little toe 


Plantar group (layer 2) 


Assists in flexing Toes 2-5 | Flexion Quadratus Medial and lateral Tendon of flexor 
toes 2-5 plantae sides of calcaneus digitorum longus 


Extends toes 2-5 at | Toes 2-5 | Extension; Lumbricals Tendons of flexor Medial side of 

the interphalangeal flexion digitorum longus proximal phalanx of 
joints; flexes the toes 2-5 

small toes at the 

metatarsophalangeal 

joints 


Plantar group (layer 3) 


Flexes big toe Big toe Flexion Flexor Lateral cuneiform; Base of proximal 
hallucis cuboid bones phalanx of big toe 
brevis 


Adducts and flexes Adduction; Adductor Bases of Base of proximal 
big toe flexion hallucis metatarsals 2—4; phalanx of big toe 
fibularis longus 
tendon sheath; 
ligament across 
metatarsophalangeal 
joints 


Flexes small toe Little toe | Flexion Flexor digiti Base of metatarsal 5; | Base of proximal 
minimi brevis | tendon sheath of phalanx of little toe 
fibularis longus 


Plantar group (layer 4) 


Abducts and flexes Middle Abduction; Dorsal Sides of metatarsals | Both sides of toe 2; 
middle toes at toes flexion; interossei for each other toe, 
metatarsophalangeal extension extensor expansion 
joints; extends over first phalanx on 
middle toes at side opposite toe 2 
interphalangeal 

joints 


Abducts toes 3-5; Small Abduction; Plantar Side of each Extensor expansion 
flexes proximal toes flexion; interossei metatarsal that faces | on first phalanx of 
phalanges and extension metatarsal 2 (absent | each toe (except to 


extends distal from metatarsal 2) 2) on side facing 
toe 2 


phalanges 


Chapter Review 


The pelvic girdle attaches the legs to the axial skeleton. The hip joint is 

where the pelvic girdle and the leg come together. The hip is joined to the 
pelvic girdle by many muscles. In the gluteal region, the psoas major and 
iliacus form the iliopsoas. The large and strong gluteus maximus, gluteus 


medius, and gluteus minimus extend and abduct the femur. Along with the 
gluteus maximus, the tensor fascia lata muscle forms the iliotibial tract. The 
lateral rotators of the femur at the hip are the piriformis, obturator internus, 
obturator externus, superior gemellus, inferior gemellus, and quadratus 
femoris. On the medial part of the thigh, the adductor longus, adductor 
brevis, and adductor magnus adduct the thigh and medially rotate it. The 
pectineus muscle adducts and flexes the femur at the hip. 


The thigh muscles that move the femur, tibia, and fibula are divided into 
medial, anterior, and posterior compartments. The medial compartment 
includes the adductors, pectineus, and the gracilis. The anterior 
compartment comprises the quadriceps femoris, quadriceps tendon, patellar 
ligament, and the sartorius. The quadriceps femoris is made of four 
muscles: the rectus femoris, the vastus lateralis, the vastus medius, and the 
vastus intermedius, which together extend the knee. The posterior 
compartment of the thigh includes the hamstrings: the biceps femoris, 
semitendinosus, and the semimembranosus, which all flex the knee. 


The muscles of the leg that move the foot and toes are divided into anterior, 
lateral, superficial- and deep-posterior compartments. The anterior 
compartment includes the tibialis anterior, the extensor hallucis longus, the 
extensor digitorum longus, and the fibularis (peroneus) tertius. The lateral 
compartment houses the fibularis (peroneus) longus and the fibularis 
(peroneus) brevis. The superficial posterior compartment has the 
gastrocnemius, soleus, and plantaris; and the deep posterior compartment 
has the popliteus, tibialis posterior, flexor digitorum longus, and flexor 
hallucis longus. 


Review Questions 


Exercise: 


Problem: 


The large muscle group that attaches the leg to the pelvic girdle and 
produces extension of the hip joint is the group. 


a. gluteal 


b. obturator 
c. adductor 
d. abductor 


Solution: 


A 
Exercise: 


Problem: 
Which muscle produces movement that allows you to cross your legs? 


a. the gluteus maximus 
b. the piriformis 

c. the gracilis 

d. the sartorius 


Solution: 


D 


Exercise: 


Problem: What is the largest muscle in the lower leg? 


a. soleus 

b. gastrocnemius 
c. tibialis anterior 
d. tibialis posterior 


Solution: 


B 


Exercise: 


Problem: 


The vastus intermedius muscle is deep to which of the following 
muscles? 


a. biceps femoris 
b. rectus femoris 

c. vastus medialis 
d. vastus lateralis 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


Which muscles form the hamstrings? How do they function together? 


Solution: 


The biceps femoris, semimembranosus, and semitendinosus form the 
hamstrings. The hamstrings flex the leg at the knee joint. 
Exercise: 


Problem: 
Which muscles form the quadriceps? How do they function together? 
Solution: 


The rectus femoris, vastus medialis, vastus lateralis, and vastus 
intermedius form the quadriceps. The quadriceps muscles extend the 
leg at the knee joint. 


Glossary 


adductor brevis 
muscle that adducts and medially rotates the thigh 


adductor longus 
muscle that adducts, medially rotates, and flexes the thigh 


adductor magnus 
muscle with an anterior fascicle that adducts, medially rotates and 
flexes the thigh, and a posterior fascicle that assists in thigh extension 


anterior compartment of the leg 
region that includes muscles that dorsiflex the foot 


anterior compartment of the thigh 
region that includes muscles that flex the thigh and extend the leg 


biceps femoris 
hamstring muscle 


calcaneal tendon 
(also, Achilles tendon) strong tendon that inserts into the calcaneal 
bone of the ankle 


dorsal group 
region that includes the extensor digitorum brevis 


extensor digitorum brevis 
muscle that extends the toes 


extensor digitorum longus 
muscle that is lateral to the tibialis anterior 


extensor hallucis longus 
muscle that is partly deep to the tibialis anterior and extensor 
digitorum longus 


femoral triangle 
region formed at the junction between the hip and the leg and includes 
the pectineus, femoral nerve, femoral artery, femoral vein, and deep 
inguinal lymph nodes 


fibularis brevis 
(also, peroneus brevis) muscle that plantar flexes the foot at the ankle 
and everts it at the intertarsal joints 


fibularis longus 
(also, peroneus longus) muscle that plantar flexes the foot at the ankle 
and everts it at the intertarsal joints 


fibularis tertius 
small muscle that is associated with the extensor digitorum longus 


flexor digitorum longus 
muscle that flexes the four small toes 


flexor hallucis longus 
muscle that flexes the big toe 


gastrocnemius 
most superficial muscle of the calf 


gluteal group 
muscle group that extends, flexes, rotates, adducts, and abducts the 
femur 


gluteus maximus 
largest of the gluteus muscles that extends the femur 


gluteus medius 
muscle deep to the gluteus maximus that abducts the femur at the hip 


gluteus minimus 
smallest of the gluteal muscles and deep to the gluteus medius 


gracilis 


muscle that adducts the thigh and flexes the leg at the knee 


hamstring group 
three long muscles on the back of the leg 


iliacus 
muscle that, along with the psoas major, makes up the iliopsoas 


iliopsoas group 
muscle group consisting of iliacus and psoas major muscles, that flexes 
the thigh at the hip, rotates it laterally, and flexes the trunk of the body 
onto the hip 


iliotibial tract 
muscle that inserts onto the tibia; made up of the gluteus maximus and 
connective tissues of the tensor fasciae latae 


inferior extensor retinaculum 
cruciate ligament of the ankle 


inferior gemellus 
muscle deep to the gluteus maximus on the lateral surface of the thigh 
that laterally rotates the femur at the hip 


lateral compartment of the leg 
region that includes the fibularis (peroneus) longus and the fibularis 
(peroneus) brevis and their associated blood vessels and nerves 


medial compartment of the thigh 
a region that includes the adductor longus, adductor brevis, adductor 
magnus, pectineus, gracilis, and their associated blood vessels and 
nerves 


obturator externus 
muscle deep to the gluteus maximus on the lateral surface of the thigh 
that laterally rotates the femur at the hip 


obturator internus 


muscle deep to the gluteus maximus on the lateral surface of the thigh 
that laterally rotates the femur at the hip 


patellar ligament 
extension of the quadriceps tendon below the patella 


pectineus 
muscle that abducts and flexes the femur at the hip 


pelvic girdle 
hips, a foundation for the lower limb 


piriformis 
muscle deep to the gluteus maximus on the lateral surface of the thigh 
that laterally rotates the femur at the hip 


plantar aponeurosis 
muscle that supports the longitudinal arch of the foot 


plantar group 
four-layered group of intrinsic foot muscles 


plantaris 
muscle that runs obliquely between the gastrocnemius and the soleus 


popliteal fossa 
diamond-shaped space at the back of the knee 


popliteus 
muscle that flexes the leg at the knee and creates the floor of the 
popliteal fossa 


posterior compartment of the leg 
region that includes the superficial gastrocnemius, soleus, and 
plantaris, and the deep popliteus, flexor digitorum longus, flexor 
hallucis longus, and tibialis posterior 


posterior compartment of the thigh 
region that includes muscles that flex the leg and extend the thigh 


psoas major 
muscle that, along with the iliacus, makes up the iliopsoas 


quadratus femoris 
muscle deep to the gluteus maximus on the lateral surface of the thigh 
that laterally rotates the femur at the hip 


quadriceps femoris group 
four muscles, that extend and stabilize the knee 


quadriceps tendon 
(also, patellar tendon) tendon common to all four quadriceps muscles, 
inserts into the patella 


rectus femoris 
quadricep muscle on the anterior aspect of the thigh 


Sartorius 
band-like muscle that flexes, abducts, and laterally rotates the leg at 
the hip 


semimembranosus 
hamstring muscle 


semitendinosus 
hamstring muscle 


soleus 
wide, flat muscle deep to the gastrocnemius 


superior extensor retinaculum 
transverse ligament of the ankle 


superior gemellus 
muscle deep to the gluteus maximus on the lateral surface of the thigh 
that laterally rotates the femur at the hip 


tensor fascia lata 
muscle that flexes and abducts the thigh 


tibialis anterior 
muscle located on the lateral surface of the tibia 


tibialis posterior 
muscle that plantar flexes and inverts the foot 


vastus intermedius 
quadricep muscle that is between the vastus lateralis and vastus 
medialis and is deep to the rectus femoris 


vastus lateralis 
quadricep muscle on the lateral aspect of the thigh 


vastus medialis 
quadricep muscle on the medial aspect of the thigh 


Heart Anatomy 
By the end of this section, you will be able to: 


e Describe the location and position of the heart within the body cavity 

e Describe the internal and external anatomy of the heart 

e Identify the tissue layers of the heart 

e Relate the structure of the heart to its function as a pump 

e Compare systemic circulation to pulmonary circulation 

e Identify the veins and arteries of the coronary circulation system 

e Trace the pathway of oxygenated and deoxygenated blood thorough 
the chambers of the heart 


The vital importance of the heart is obvious. If one assumes an average rate 
of contraction of 75 contractions per minute, a human heart would contract 
approximately 108,000 times in one day, more than 39 million times in one 
year, and nearly 3 billion times during a 75-year lifespan. Each of the major 
pumping chambers of the heart ejects approximately 70 mL blood per 
contraction in a resting adult. This would be equal to 5.25 liters of fluid per 
minute and approximately 14,000 liters per day. Over one year, that would 
equal 10,000,000 liters or 2.6 million gallons of blood sent through roughly 
60,000 miles of vessels. In order to understand how that happens, it is 
necessary to understand the anatomy and physiology of the heart. 


Location of the Heart 


The human heart is located within the thoracic cavity, medially between the 
lungs in the space known as the mediastinum. [link] shows the position of 
the heart within the thoracic cavity. Within the mediastinum, the heart is 
separated from the other mediastinal structures by a tough membrane 
known as the pericardium, or pericardial sac, and sits in its own space 
called the pericardial cavity. The dorsal surface of the heart lies near the 
bodies of the vertebrae, and its anterior surface sits deep to the sternum and 
costal cartilages. The great veins, the superior and inferior venae cavae, and 
the great arteries, the aorta and pulmonary trunk, are attached to the 
superior surface of the heart, called the base. The base of the heart is 
located at the level of the third costal cartilage, as seen in [link]. The 
inferior tip of the heart, the apex, lies just to the left of the sternum between 


the junction of the fourth and fifth ribs near their articulation with the costal 
cartilages. The right side of the heart is deflected anteriorly, and the left side 
is deflected posteriorly. It is important to remember the position and 
orientation of the heart when placing a stethoscope on the chest of a patient 
and listening for heart sounds, and also when looking at images taken from 
a midsagittal perspective. The slight deviation of the apex to the left is 
reflected in a depression in the medial surface of the inferior lobe of the left 
lung, called the cardiac notch. 

Position of the Heart in the Thorax 


Thoracic 
aorta 


Sagittal view 


Mediastinum 


; Arch of aorta 
Superior vena cava 


Right lung Pulmonary trunk 


Right auricle Left auricle 


Right atrium Left lung 


Right ventricle Left ventricle 


Ribs (cut) Pericardial cavity 
Ditiimwiesien 


Apex of heart 


Edge of parietal Edge of parietal 
pleura (cut) pericardium (cut) 


The heart is located within the thoracic cavity, medially 
between the lungs in the mediastinum. It is about the size 
of a fist, is broad at the top, and tapers toward the base. 


Note: 

Everyday Connection 

CPR 

The position of the heart in the torso between the vertebrae and sternum 
(see [link] for the position of the heart within the thorax) allows for 
individuals to apply an emergency technique known as cardiopulmonary 
resuscitation (CPR) if the heart of a patient should stop. By applying 
pressure with the flat portion of one hand on the sternum in the area 
between the line at T4 and T9 ((link]), it is possible to manually compress 
the blood within the heart enough to push some of the blood within it into 
the pulmonary and systemic circuits. This is particularly critical for the 
brain, as irreversible damage and death of neurons occur within minutes of 
loss of blood flow. Current standards call for compression of the chest at 
least 5 cm deep and at a rate of 100 compressions per minute, a rate equal 
to the beat in “Staying Alive,” recorded in 1977 by the Bee Gees. If you 
are unfamiliar with this song, a version is available on www.youtube.com. 
At this stage, the emphasis is on performing high-quality chest 
compressions, rather than providing artificial respiration. CPR is generally 
performed until the patient regains spontaneous contraction or is declared 
dead by an experienced healthcare professional. 

When performed by untrained or overzealous individuals, CPR can result 
in broken ribs or a broken sternum, and can inflict additional severe 
damage on the patient. It is also possible, if the hands are placed too low 
on the sternum, to manually drive the xiphoid process into the liver, a 
consequence that may prove fatal for the patient. Proper training is 
essential. This proven life-sustaining technique is so valuable that virtually 
all medical personnel as well as concerned members of the public should 
be certified and routinely recertified in its application. CPR courses are 
offered at a variety of locations, including colleges, hospitals, the 
American Red Cross, and some commercial companies. They normally 
include practice of the compression technique on a mannequin. 

CPR Technique 


If the heart should stop, CPR can maintain 
the flow of blood until the heart resumes 
beating. By applying pressure to the 
sternum, the blood within the heart will be 
squeezed out of the heart and into the 
circulation. Proper positioning of the 
hands on the sternum to perform CPR 
would be between the lines at T4 and T9. 


Note: 


meee OPENStAX COLLEGE 
it ‘s 
. L 
OS 


Visit the American Heart Association website to help locate a course near 
your home in the United States. There are also many other national and 


regional heart associations that offer the same service, depending upon the 
location. 


Shape and Size of the Heart 


The shape of the heart is similar to a pinecone, rather broad at the superior 
surface and tapering to the apex (see [link]). A typical heart is 
approximately the size of your fist: 12 cm (5 in) in length, 8 cm (3.5 in) 
wide, and 6 cm (2.5 in) in thickness. Given the size difference between 
most members of the sexes, the weight of a female heart is approximately 
250-300 grams (9 to 11 ounces), and the weight of a male heart is 
approximately 300-350 grams (11 to 12 ounces). The heart of a well- 
trained athlete, especially one specializing in aerobic sports, can be 
considerably larger than this. Cardiac muscle responds to exercise in a 
manner similar to that of skeletal muscle. That is, exercise results in the 
addition of protein myofilaments that increase the size of the individual 
cells without increasing their numbers, a concept called hypertrophy. Hearts 
of athletes can pump blood more effectively at lower rates than those of 
nonathletes. Enlarged hearts are not always a result of exercise; they can 
result from pathologies, such as hypertrophic cardiomyopathy. The cause 
of an abnormally enlarged heart muscle is unknown, but the condition is 
often undiagnosed and can cause sudden death in apparently otherwise 
healthy young people. 


Chambers and Circulation through the Heart 


The human heart consists of four chambers: The left side and the right side 
each have one atrium and one ventricle. Each of the upper chambers, the 
right atrium (plural = atria) and the left atrium, acts as a receiving chamber 
and contracts to push blood into the lower chambers, the right ventricle and 
the left ventricle. The ventricles serve as the primary pumping chambers of 
the heart, propelling blood to the lungs or to the rest of the body. 


There are two distinct but linked circuits in the human circulation called the 
pulmonary and systemic circuits. Although both circuits transport blood and 


everything it carries, we can initially view the circuits from the point of 
view of gases. The pulmonary circuit transports blood to and from the 
lungs, where it picks up oxygen and delivers carbon dioxide for exhalation. 
The systemic circuit transports oxygenated blood to virtually all of the 
tissues of the body and returns relatively deoxygenated blood and carbon 
dioxide to the heart to be sent back to the pulmonary circulation. 


The right ventricle pumps deoxygenated blood into the pulmonary trunk, 
which leads toward the lungs and bifurcates into the left and right 
pulmonary arteries. These vessels in turn branch many times before 
reaching the pulmonary capillaries, where gas exchange occurs: Carbon 
dioxide exits the blood and oxygen enters. The pulmonary trunk arteries 
and their branches are the only arteries in the post-natal body that carry 
relatively deoxygenated blood. Highly oxygenated blood returning from the 
pulmonary capillaries in the lungs passes through a series of vessels that 
join together to form the pulmonary veins—the only post-natal veins in the 
body that carry highly oxygenated blood. The pulmonary veins conduct 
blood into the left atrium, which pumps the blood into the left ventricle, 
which in turn pumps oxygenated blood into the aorta and on to the many 
branches of the systemic circuit. Eventually, these vessels will lead to the 
systemic capillaries, where exchange with the tissue fluid and cells of the 
body occurs. In this case, oxygen and nutrients exit the systemic capillaries 
to be used by the cells in their metabolic processes, and carbon dioxide and 
waste products will enter the blood. 


The blood exiting the systemic capillaries is lower in oxygen concentration 
than when it entered. The capillaries will ultimately unite to form venules, 
joining to form ever-larger veins, eventually flowing into the two major 
systemic veins, the superior vena cava and the inferior vena cava, which 
return blood to the right atrium. The blood in the superior and inferior 
venae cavae flows into the right atrium, which pumps blood into the right 
ventricle. This process of blood circulation continues as long as the 
individual remains alive. Understanding the flow of blood through the 
pulmonary and systemic circuits is critical to all health professions ({link]). 
Dual System of the Human Blood Circulation 


Aorta 


Left pulmonary 
Right pulmonary arteries 


arteries 
Pulmonary trunk 


Left atrium 
Right pulmonary 
veins Left pulmonary 
veins 
Pulmonary 
semilunar 
valve 


Aortic semilunar 
valve 


Mitral valve 
Right atrium 
Tricuspid valve Left ventricle 


Right ventricle 


Systemic Systemic 
veins from capillaries 
upper body of upper 


body 


Systemic 
arteries to 


Pulmona 
es upper body 


capillaries 
in lungs 

Pulmonary 
Right atrium trunk 
Left atrium 
Right 


ventricle Left 

\ ventricle 
Systemic 
veins from 


Systemic 


lower body arteries to 
lower body 

Systemic 

capillaries 


of lower body 


Blood flows from the right atrium to the right ventricle, 
where it is pumped into the pulmonary circuit. The blood in 
the pulmonary artery branches is low in oxygen but 
relatively high in carbon dioxide. Gas exchange occurs in 
the pulmonary capillaries (oxygen into the blood, carbon 
dioxide out), and blood high in oxygen and low in carbon 
dioxide is returned to the left atrium. From here, blood 
enters the left ventricle, which pumps it into the systemic 
circuit. Following exchange in the systemic capillaries 
(oxygen and nutrients out of the capillaries and carbon 


dioxide and wastes in), blood returns to the right atrium and 
the cycle is repeated. 


Membranes, Surface Features, and Layers 


Our exploration of more in-depth heart structures begins by examining the 
membrane that surrounds the heart, the prominent surface features of the 
heart, and the layers that form the wall of the heart. Each of these 
components plays its own unique role in terms of function. 


Membranes 


The membrane that directly surrounds the heart and defines the pericardial 
cavity is called the pericardium or pericardial sac. It also surrounds the 
“roots” of the major vessels, or the areas of closest proximity to the heart. 
The pericardium, which literally translates as “around the heart,” consists of 
two distinct sublayers: the sturdy outer fibrous pericardium and the inner 
serous pericardium. The fibrous pericardium is made of tough, dense 
connective tissue that protects the heart and maintains its position in the 
thorax. The more delicate serous pericardium consists of two layers: the 
parietal pericardium, which is fused to the fibrous pericardium, and an inner 
visceral pericardium, or epicardium, which is fused to the heart and is part 
of the heart wall. The pericardial cavity, filled with lubricating serous fluid, 
lies between the epicardium and the pericardium. 


In most organs within the body, visceral serous membranes such as the 
epicardium are microscopic. However, in the case of the heart, it is not a 
microscopic layer but rather a macroscopic layer, consisting of a simple 
squamous epithelium called a mesothelium, reinforced with loose, 
irregular, or areolar connective tissue that attaches to the pericardium. This 
mesothelium secretes the lubricating serous fluid that fills the pericardial 
cavity and reduces friction as the heart contracts. [link] illustrates the 
pericardial membrane and the layers of the heart. 

Pericardial Membranes and Layers of the Heart Wall 


Pericardial cavity 


Endocardium 


Fibrous pericardium 
Myocardium 


Parietal layer of serous 
pericardium 


Epicardium (viceral layer 
of serous pericardium) 


The pericardial membrane that surrounds the heart 
consists of three layers and the pericardial cavity. The 
heart wall also consists of three layers. The pericardial 

membrane and the heart wall share the epicardium. 


Note: 

Disorders of the... 

Heart: Cardiac Tamponade 

If excess fluid builds within the pericardial space, it can lead to a condition 
called cardiac tamponade, or pericardial tamponade. With each contraction 
of the heart, more fluid—in most instances, blood—accumulates within the 
pericardial cavity. In order to fill with blood for the next contraction, the 
heart must relax. However, the excess fluid in the pericardial cavity puts 
pressure on the heart and prevents full relaxation, so the chambers within 
the heart contain slightly less blood as they begin each heart cycle. Over 
time, less and less blood is ejected from the heart. If the fluid builds up 
slowly, as in hypothyroidism, the pericardial cavity may be able to expand 
gradually to accommodate this extra volume. Some cases of fluid in excess 


of one liter within the pericardial cavity have been reported. Rapid 
accumulation of as little as 100 mL of fluid following trauma may trigger 
cardiac tamponade. Other common causes include myocardial rupture, 
pericarditis, cancer, or even cardiac surgery. Removal of this excess fluid 
requires insertion of drainage tubes into the pericardial cavity. Premature 
removal of these drainage tubes, for example, following cardiac surgery, or 
clot formation within these tubes are causes of this condition. Untreated, 
cardiac tamponade can lead to death. 


Surface Features of the Heart 


Inside the pericardium, the surface features of the heart are visible, 
including the four chambers. There is a superficial leaf-like extension of the 
atria near the superior surface of the heart, one on each side, called an 
auricle—a name that means “ear like”—because its shape resembles the 
external ear of a human ([link]). Auricles are relatively thin-walled 
structures that can fill with blood and empty into the atria or upper 
chambers of the heart. You may also hear them referred to as atrial 
appendages. Also prominent is a series of fat-filled grooves, each of which 
is known as a sulcus (plural = sulci), along the superior surfaces of the 
heart. Major coronary blood vessels are located in these sulci. The deep 
coronary sulcus is located between the atria and ventricles. Located 
between the left and right ventricles are two additional sulci that are not as 
deep as the coronary sulcus. The anterior interventricular sulcus is 
visible on the anterior surface of the heart, whereas the posterior 
interventricular sulcus is visible on the posterior surface of the heart. 
[link] illustrates anterior and posterior views of the surface of the heart. 
External Anatomy of the Heart 


Brachiocephalic trunk Left common carotid artery 
Left subclavian artery 


Aortic arch 
Ligamentum arteriosum 


Left pulmonary artery 


Superior vena cava 


Right pulmonary 
artery 


Ascending aorta 
Pulmonary trunk 


 F 
Right pulmonary << é 


veins 


Left pulmonary veins 


Auricle of left atrium 
Anterior view 


Right atrium Circumflex artery 


Right coronary artery Left coronary artery 


Anterior cardiac vein 
: ; Left ventricle 
Right ventricle 


Great cardiac vein 
Anterior interventricular artery 


Right marginal artery 
Small cardiac vein 
Inferior vena cava 
Apex 


Superior vena cava 
Right pul t 
Left pulmonary artery ignt pulmonary artery 


. Right pulmonary veins 
Left pulmonary veins 


Auricle of left atrium Right atrium 


Left atrium Inferior vena cava 


Circumflex branch Coronary sinus 


Posterior view of left coronary artery Small cardiac vein 


Great cardiac vein Right coronary artery 


Posterior vein of 


left ventricle Posterior interventricular artery 


Middle cardiac vein 
Left ventricle 


Right ventricle 


Inside the pericardium, the surface features of the heart are 
visible. 


Layers 


The wall of the heart is composed of three layers of unequal thickness. 
From superficial to deep, these are the epicardium, the myocardium, and the 
endocardium (see [link]). The outermost layer of the wall of the heart is 
also the innermost layer of the pericardium, the epicardium, or the visceral 
pericardium discussed earlier. 


The middle and thickest layer is the myocardium, made largely of cardiac 
muscle cells. It is built upon a framework of collagenous fibers, plus the 
blood vessels that supply the myocardium and the nerve fibers that help 
regulate the heart. It is the contraction of the myocardium that pumps blood 
through the heart and into the major arteries. The muscle pattern is elegant 
and complex, as the muscle cells swirl and spiral around the chambers of 
the heart. They form a figure 8 pattern around the atria and around the bases 
of the great vessels. Deeper ventricular muscles also form a figure 8 around 
the two ventricles and proceed toward the apex. More superficial layers of 
ventricular muscle wrap around both ventricles. This complex swirling 
pattern allows the heart to pump blood more effectively than a simple linear 
pattern would. [link] illustrates the arrangement of muscle cells. 

Heart Musculature 


Atrial 
musculature 


Ventricular 
musculature 


The swirling pattern of 
cardiac muscle tissue 
contributes significantly to 
the heart’s ability to pump 
blood effectively. 


Although the ventricles on the right and left sides pump the same amount of 
blood per contraction, the muscle of the left ventricle is much thicker and 
better developed than that of the right ventricle. In order to overcome the 


high resistance required to pump blood into the long systemic circuit, the 
left ventricle must generate a great amount of pressure. The right ventricle 
does not need to generate as much pressure, since the pulmonary circuit is 
shorter and provides less resistance. [link] illustrates the differences in 
muscular thickness needed for each of the ventricles. 

Differences in Ventricular Muscle Thickness 


Left ventricle 


Right ventricle 


Relaxed Contracted 


The myocardium in the left ventricle is significantly 
thicker than that of the right ventricle. Both ventricles 
pump the same amount of blood, but the left ventricle 

must generate a much greater pressure to overcome 

greater resistance in the systemic circuit. The 
ventricles are shown in both relaxed and contracting 
states. Note the differences in the relative size of the 
lumens, the region inside each ventricle where the 
blood is contained. 


The innermost layer of the heart wall, the endocardium, is joined to the 
myocardium with a thin layer of connective tissue. The endocardium lines 
the chambers where the blood circulates and covers the heart valves. It is 
made of simple squamous epithelium called endothelium, which is 
continuous with the endothelial lining of the blood vessels (see [link]). 


Once regarded as a simple lining layer, recent evidence indicates that the 
endothelium of the endocardium and the coronary capillaries may play 
active roles in regulating the contraction of the muscle within the 
myocardium. The endothelium may also regulate the growth patterns of the 
cardiac muscle cells throughout life, and the endothelins it secretes create 
an environment in the surrounding tissue fluids that regulates ionic 
concentrations and states of contractility. Endothelins are potent 
vasoconstrictors and, in a normal individual, establish a homeostatic 
balance with other vasoconstrictors and vasodilators. 


Internal Structure of the Heart 


Recall that the heart’s contraction cycle follows a dual pattern of circulation 
—the pulmonary and systemic circuits—because of the pairs of chambers 
that pump blood into the circulation. In order to develop a more precise 
understanding of cardiac function, it is first necessary to explore the internal 
anatomical structures in more detail. 


Septa of the Heart 


The word septum is derived from the Latin for “something that encloses;” 
in this case, a septum (plural = septa) refers to a wall or partition that 
divides the heart into chambers. The septa are physical extensions of the 
myocardium lined with endocardium. Located between the two atria is the 
interatrial septum. Normally in an adult heart, the interatrial septum bears 
an oval-shaped depression known as the fossa ovalis, a remnant of an 
opening in the fetal heart known as the foramen ovale. The foramen ovale 
allowed blood in the fetal heart to pass directly from the right atrium to the 
left atrium, allowing some blood to bypass the pulmonary circuit. Within 
seconds after birth, a flap of tissue known as the septum primum that 
previously acted as a valve closes the foramen ovale and establishes the 
typical cardiac circulation pattern. 


Between the two ventricles is a second septum known as the 
interventricular septum. Unlike the interatrial septum, the interventricular 


septum is normally intact after its formation during fetal development. It is 
substantially thicker than the interatrial septum, since the ventricles 
generate far greater pressure when they contract. 


The septum between the atria and ventricles is known as the 
atrioventricular septum. It is marked by the presence of four openings 
that allow blood to move from the atria into the ventricles and from the 
ventricles into the pulmonary trunk and aorta. Located in each of these 
openings between the atria and ventricles is a valve, a specialized structure 
that ensures one-way flow of blood. The valves between the atria and 
ventricles are known generically as atrioventricular valves. The valves at 
the openings that lead to the pulmonary trunk and aorta are known 
generically as semilunar valves. The interventricular septum is visible in 
[link]. In this figure, the atrioventricular septum has been removed to better 
show the bicupid and tricuspid valves; the interatrial septum is not visible, 
since its location is covered by the aorta and pulmonary trunk. Since these 
openings and valves structurally weaken the atrioventricular septum, the 
remaining tissue is heavily reinforced with dense connective tissue called 
the cardiac skeleton, or skeleton of the heart. It includes four rings that 
surround the openings between the atria and ventricles, and the openings to 
the pulmonary trunk and aorta, and serve as the point of attachment for the 
heart valves. The cardiac skeleton also provides an important boundary in 
the heart electrical conduction system. 

Internal Structures of the Heart 


Superior vena cava Left pulmonary artery 


Right pulmonary artery peiaidum 


Pulmonary trunk Left pulmonary veins 


Right pulmonary 
veins Mitral (bicuspid) valve 


Right atrium 
Aortic valve 


Fossa ovalis 
Tricuspid valve Pulmonary valve 


Right ventricle Left ventricle 


Chordae tendineae Papillary muscle 
Interventricular septum 
Epicardium 
Myocardium 
Endocardium 


Trabeculae carneae 


Moderator band 


Inferior vena cava 


Anterior view 


This anterior view of the heart shows the four 
chambers, the major vessels and their early branches, 
as well as the valves. The presence of the pulmonary 
trunk and aorta covers the interatrial septum, and the 

atrioventricular septum is cut away to show the 
atrioventricular valves. 


Note: 

Disorders of the... 

Heart: Heart Defects 

One very common form of interatrial septum pathology is patent foramen 
ovale, which occurs when the septum primum does not close at birth, and 
the fossa ovalis is unable to fuse. The word patent is from the Latin root 
patens for “open.” It may be benign or asymptomatic, perhaps never being 
diagnosed, or in extreme cases, it may require surgical repair to close the 
opening permanently. As much as 20—25 percent of the general population 
may have a patent foramen ovale, but fortunately, most have the benign, 


asymptomatic version. Patent foramen ovale is normally detected by 
auscultation of a heart murmur (an abnormal heart sound) and confirmed 
by imaging with an echocardiogram. Despite its prevalence in the general 
population, the causes of patent ovale are unknown, and there are no 
known risk factors. In nonlife-threatening cases, it is better to monitor the 
condition than to risk heart surgery to repair and seal the opening. 
Coarctation of the aorta is a congenital abnormal narrowing of the aorta 
that is normally located at the insertion of the ligamentum arteriosum, the 
remnant of the fetal shunt called the ductus arteriosus. If severe, this 
condition drastically restricts blood flow through the primary systemic 
artery, which is life threatening. In some individuals, the condition may be 
fairly benign and not detected until later in life. Detectable symptoms in an 
infant include difficulty breathing, poor appetite, trouble feeding, or failure 
to thrive. In older individuals, symptoms include dizziness, fainting, 
shortness of breath, chest pain, fatigue, headache, and nosebleeds. 
Treatment involves surgery to resect (remove) the affected region or 
angioplasty to open the abnormally narrow passageway. Studies have 
shown that the earlier the surgery is performed, the better the chance of 
survival. 

A patent ductus arteriosus is a congenital condition in which the ductus 
arteriosus fails to close. The condition may range from severe to benign. 
Failure of the ductus arteriosus to close results in blood flowing from the 
higher pressure aorta into the lower pressure pulmonary trunk. This 
additional fluid moving toward the lungs increases pulmonary pressure and 
makes respiration difficult. Symptoms include shortness of breath 
(dyspnea), tachycardia, enlarged heart, a widened pulse pressure, and poor 
weight gain in infants. Treatments include surgical closure (ligation), 
manual closure using platinum coils or specialized mesh inserted via the 
femoral artery or vein, or nonsteroidal anti-inflammatory drugs to block 
the synthesis of prostaglandin E2, which maintains the vessel in an open 
position. If untreated, the condition can result in congestive heart failure. 
Septal defects are not uncommon in individuals and may be congenital or 
caused by various disease processes. Tetralogy of Fallot is a congenital 
condition that may also occur from exposure to unknown environmental 
factors; it occurs when there is an opening in the interventricular septum 
caused by blockage of the pulmonary trunk, normally at the pulmonary 
semilunar valve. This allows blood that is relatively low in oxygen from 


the right ventricle to flow into the left ventricle and mix with the blood that 
is relatively high in oxygen. Symptoms include a distinct heart murmur, 
low blood oxygen percent saturation, dyspnea or difficulty in breathing, 
polycythemia, broadening (clubbing) of the fingers and toes, and in 
children, difficulty in feeding or failure to grow and develop. It is the most 
common cause of cyanosis following birth. The term “tetralogy” is derived 
from the four components of the condition, although only three may be 
present in an individual patient: pulmonary infundibular stenosis (rigidity 
of the pulmonary valve), overriding aorta (the aorta is shifted above both 
ventricles), ventricular septal defect (opening), and right ventricular 
hypertrophy (enlargement of the right ventricle). Other heart defects may 
also accompany this condition, which is typically confirmed by 
echocardiography imaging. Tetralogy of Fallot occurs in approximately 
AO0 out of one million live births. Normal treatment involves extensive 
surgical repair, including the use of stents to redirect blood flow and 
replacement of valves and patches to repair the septal defect, but the 
condition has a relatively high mortality. Survival rates are currently 75 
percent during the first year of life; 60 percent by 4 years of age; 30 
percent by 10 years; and 5 percent by 40 years. 

In the case of severe septal defects, including both tetralogy of Fallot and 
patent foramen ovale, failure of the heart to develop properly can lead to a 
condition commonly known as a “blue baby.” Regardless of normal skin 
pigmentation, individuals with this condition have an insufficient supply of 
oxygenated blood, which leads to cyanosis, a blue or purple coloration of 
the skin, especially when active. 

Septal defects are commonly first detected through auscultation, listening 
to the chest using a stethoscope. In this case, instead of hearing normal 
heart sounds attributed to the flow of blood and closing of heart valves, 
unusual heart sounds may be detected. This is often followed by medical 
imaging to confirm or rule out a diagnosis. In many cases, treatment may 
not be needed. Some common congenital heart defects are illustrated in 
[link]. 

Congenital Heart Defects 


Narrow segment 
Ha of aorta 
Foramen ovale 
fails to close 


(a) Patent foramen ovale (b) Coarctation of the aorta 


Ductus arteriosus 

remains open 
Stenosed 
pulmonary 
semilunar valve 


(c) Patent ductus arteriosus (d) Tetralogy of Fallot 


Aorta emerges 
from both 
ventricles 


Interventricular 
septal defect 


Enlarged right 
ventricle 


(a) A patent foramen ovale defect is an abnormal opening 
in the interatrial septum, or more commonly, a failure of the 
foramen ovale to close. (b) Coarctation of the aorta is an 
abnormal narrowing of the aorta. (c) A patent ductus 
arteriosus is the failure of the ductus arteriosus to close. (d) 
Tetralogy of Fallot includes an abnormal opening in the 
interventricular septum. 


Right Atrium 


The right atrium serves as the receiving chamber for blood returning to the 
heart from the systemic circulation. The two major systemic veins, the 
superior and inferior venae cavae, and the large coronary vein called the 
coronary sinus that drains the heart myocardium empty into the right 
atrium. The superior vena cava drains blood from regions superior to the 
diaphragm: the head, neck, upper limbs, and the thoracic region. It empties 
into the superior and posterior portions of the right atrium. The inferior 
vena cava drains blood from areas inferior to the diaphragm: the lower 
limbs and abdominopelvic region of the body. It, too, empties into the 


posterior portion of the atria, but inferior to the opening of the superior vena 
cava. Immediately superior and slightly medial to the opening of the 
inferior vena cava on the posterior surface of the atrium is the opening of 
the coronary sinus. This thin-walled vessel drains most of the coronary 
veins that return systemic blood from the heart. The majority of the internal 
heart structures discussed in this and subsequent sections are illustrated in 
[link]. 


While the bulk of the internal surface of the right atrium is smooth, the 
depression of the fossa ovalis is medial, and the anterior surface 
demonstrates prominent ridges of muscle called the pectinate muscles. The 
right auricle also has pectinate muscles. The left atrium does not have 
pectinate muscles except in the auricle. 


The atria receive venous blood on a nearly continuous basis, preventing 
venous flow from stopping while the ventricles are contracting. While most 
ventricular filling occurs while the atria are relaxed, they do demonstrate a 
contractile phase and actively pump blood into the ventricles just prior to 
ventricular contraction. The opening between the atrium and ventricle is 
guarded by the tricuspid valve. 


Right Ventricle 


The right ventricle receives blood from the right atrium through the 
tricuspid valve. Each flap of the valve is attached to strong strands of 
connective tissue, the chordae tendineae, literally “tendinous cords,” or 
sometimes more poetically referred to as “heart strings.” There are several 
chordae tendineae associated with each of the flaps. They are composed of 
approximately 80 percent collagenous fibers with the remainder consisting 
of elastic fibers and endothelium. They connect each of the flaps to a 
papillary muscle that extends from the inferior ventricular surface. There 
are three papillary muscles in the right ventricle, called the anterior, 
posterior, and septal muscles, which correspond to the three sections of the 
valves. 


When the myocardium of the ventricle contracts, pressure within the 
ventricular chamber rises. Blood, like any fluid, flows from higher pressure 
to lower pressure areas, in this case, toward the pulmonary trunk and the 
atrium. To prevent any potential backflow, the papillary muscles also 
contract, generating tension on the chordae tendineae. This prevents the 
flaps of the valves from being forced into the atria and regurgitation of the 
blood back into the atria during ventricular contraction. [link] shows 
papillary muscles and chordae tendineae attached to the tricuspid valve. 
Chordae Tendineae and Papillary Muscles 


Chordae tendineae 
Papillary muscles 


Trabeculae carneae 


In this frontal section, you can see papillary 
muscles attached to the tricuspid valve on 
the right as well as the mitral valve on the 

left via chordae tendineae. (credit: 
modification of work by “PV 
KS”/flickr.com) 


The walls of the ventricle are lined with trabeculae carneae, ridges of 
cardiac muscle covered by endocardium. In addition to these muscular 
ridges, a band of cardiac muscle, also covered by endocardium, known as 
the moderator band (see [link]) reinforces the thin walls of the right 
ventricle and plays a crucial role in cardiac conduction. It arises from the 


inferior portion of the interventricular septum and crosses the interior space 
of the right ventricle to connect with the inferior papillary muscle. 


When the right ventricle contracts, it ejects blood into the pulmonary trunk, 
which branches into the left and right pulmonary arteries that carry it to 
each lung. The superior surface of the right ventricle begins to taper as it 
approaches the pulmonary trunk. At the base of the pulmonary trunk is the 
pulmonary semilunar valve that prevents backflow from the pulmonary 
trunk. 


Left Atrium 


After exchange of gases in the pulmonary capillaries, blood returns to the 
left atrium high in oxygen via one of the four pulmonary veins. While the 
left atrium does not contain pectinate muscles, it does have an auricle that 
includes these pectinate ridges. Blood flows nearly continuously from the 
pulmonary veins back into the atrium, which acts as the receiving chamber, 
and from here through an opening into the left ventricle. Most blood flows 
passively into the heart while both the atria and ventricles are relaxed, but 
toward the end of the ventricular relaxation period, the left atrium will 
contract, pumping blood into the ventricle. This atrial contraction accounts 
for approximately 20 percent of ventricular filling. The opening between 
the left atrium and ventricle is guarded by the mitral valve. 


Left Ventricle 


Recall that, although both sides of the heart will pump the same amount of 
blood, the muscular layer is much thicker in the left ventricle compared to 
the right (see [link]). Like the right ventricle, the left also has trabeculae 
carneae, but there is no moderator band. The mitral valve is connected to 
papillary muscles via chordae tendineae. There are two papillary muscles 
on the left—the anterior and posterior—as opposed to three on the right. 


The left ventricle is the major pumping chamber for the systemic circuit; it 
ejects blood into the aorta through the aortic semilunar valve. 


Heart Valve Structure and Function 


A transverse section through the heart slightly above the level of the 
atrioventricular septum reveals all four heart valves along the same plane 
({link]). The valves ensure unidirectional blood flow through the heart. 
Between the right atrium and the right ventricle is the right 
atrioventricular valve, or tricuspid valve. It typically consists of three 
flaps, or leaflets, made of endocardium reinforced with additional 
connective tissue. The flaps are connected by chordae tendineae to the 
papillary muscles, which control the opening and closing of the valves. 
Heart Valves 


Posterior 
Tricuspid valve Bicuspid (mitral) 


Aortic valve Pulmonary valve 


Anterior 


With the atria and major vessels removed, all 
four valves are clearly visible, although it is 
difficult to distinguish the three separate cusps 
of the tricuspid valve. 


Emerging from the right ventricle at the base of the pulmonary trunk is the 
pulmonary semilunar valve, or the pulmonary valve; it is also known as 
the pulmonic valve or the right semilunar valve. The pulmonary valve is 
comprised of three small flaps of endothelium reinforced with connective 
tissue. When the ventricle relaxes, the pressure differential causes blood to 
flow back into the ventricle from the pulmonary trunk. This flow of blood 
fills the pocket-like flaps of the pulmonary valve, causing the valve to close 
and producing an audible sound. Unlike the atrioventricular valves, there 
are no papillary muscles or chordae tendineae associated with the 
pulmonary valve. 


Located at the opening between the left atrium and left ventricle is the 
mitral valve, also called the bicuspid valve or the left atrioventricular 
valve. Structurally, this valve consists of two cusps, known as the anterior 
medial cusp and the posterior medial cusp, compared to the three cusps of 
the tricuspid valve. In a clinical setting, the valve is referred to as the mitral 
valve, rather than the bicuspid valve. The two cusps of the mitral valve are 
attached by chordae tendineae to two papillary muscles that project from 
the wall of the ventricle. 


At the base of the aorta is the aortic semilunar valve, or the aortic valve, 
which prevents backflow from the aorta. It normally is composed of three 
flaps. When the ventricle relaxes and blood attempts to flow back into the 
ventricle from the aorta, blood will fill the cusps of the valve, causing it to 
close and producing an audible sound. 


In [link]a, the two atrioventricular valves are open and the two semilunar 
valves are closed. This occurs when both atria and ventricles are relaxed 
and when the atria contract to pump blood into the ventricles. [link]b shows 
a frontal view. Although only the left side of the heart is illustrated, the 
process is virtually identical on the right. 

Blood Flow from the Left Atrium to the Left Ventricle 


Posterior 


Tricuspid Bicuspid (mitral) valve 
valve 


\ Left 
side of WA . < \ side of 


heart 


valve 
(closed) 


Anterior (closed) 


(a) Mitral 
valve 
(open) 


Aortic valve 
(Closed) 


Chordae 
tendineae 
(loose) 


Papillary 
muscle 
(relaxed) 


(b) 


(a) A transverse section through the heart illustrates 
the four heart valves. The two atrioventricular valves 
are open; the two semilunar valves are closed. The 
atria and vessels have been removed. (b) A frontal 
section through the heart illustrates blood flow 
through the mitral valve. When the mitral valve is 
open, it allows blood to move from the left atrium to 
the left ventricle. The aortic semilunar valve is closed 
to prevent backflow of blood from the aorta to the left 
ventricle. 


[link]a shows the atrioventricular valves closed while the two semilunar 
valves are open. This occurs when the ventricles contract to eject blood into 
the pulmonary trunk and aorta. Closure of the two atrioventricular valves 


prevents blood from being forced back into the atria. This stage can be seen 
from a frontal view in [link |b. 
Blood Flow from the Left Ventricle into the Great Vessels 


Posterior 


Tricuspid Bicuspid (mitral) valve 
valve - 5 ——— Jf (closed) 
\.. {~ 


Aortic 
valve 
Pulmonary 
(open) valve 
Anterior open) 
(a) Mitral 
valve 
(closed) 


Aortic valve 
(open) 
Chordae 
tendineae 
(tight) 
Papillary 
muscle 
(contracted) 


(b) 


(a) A transverse section through the heart illustrates 
the four heart valves during ventricular contraction. 
The two atrioventricular valves are closed, but the two 
semilunar valves are open. The atria and vessels have 
been removed. (b) A frontal view shows the closed 
mitral (bicuspid) valve that prevents backflow of 
blood into the left atrium. The aortic semilunar valve 
is open to allow blood to be ejected into the aorta. 


When the ventricles begin to contract, pressure within the ventricles rises 
and blood flows toward the area of lowest pressure, which is initially in the 


atria. This backflow causes the cusps of the tricuspid and mitral (bicuspid) 
valves to close. These valves are tied down to the papillary muscles by 
chordae tendineae. During the relaxation phase of the cardiac cycle, the 
papillary muscles are also relaxed and the tension on the chordae tendineae 
is slight (see [link]b). However, as the myocardium of the ventricle 
contracts, so do the papillary muscles. This creates tension on the chordae 
tendineae (see [link]b), helping to hold the cusps of the atrioventricular 
valves in place and preventing them from being blown back into the atria. 


The aortic and pulmonary semilunar valves lack the chordae tendineae and 
papillary muscles associated with the atrioventricular valves. Instead, they 
consist of pocket-like folds of endocardium reinforced with additional 
connective tissue. When the ventricles relax and the change in pressure 
forces the blood toward the ventricles, the blood presses against these cusps 
and seals the openings. 


Note: 


— 
meee OPENSTAX COLLEGE 


F 


Bear es 


Visit this site to observe an echocardiogram of actual heart valves opening 
and closing. Although much of the heart has been “removed” from this gif 
loop so the chordae tendineae are not visible, why is their presence more 
critical for the atrioventricular valves (tricuspid and mitral) than the 
semilunar (aortic and pulmonary) valves? 


Note: 
Disorders of the... 
Heart Valves 


When heart valves do not function properly, they are often described as 
incompetent and result in valvular heart disease, which can range from 
benign to lethal. Some of these conditions are congenital, that is, the 
individual was born with the defect, whereas others may be attributed to 
disease processes or trauma. Some malfunctions are treated with 
medications, others require surgery, and still others may be mild enough 
that the condition is merely monitored since treatment might trigger more 
serious Consequences. 

Valvular disorders are often caused by carditis, or inflammation of the 
heart. One common trigger for this inflammation is rheumatic fever, or 
scarlet fever, an autoimmune response to the presence of a bacterium, 
Streptococcus pyogenes, normally a disease of childhood. 

While any of the heart valves may be involved in valve disorders, mitral 
regurgitation is the most common, detected in approximately 2 percent of 
the population, and the pulmonary semilunar valve is the least frequently 
involved. When a valve malfunctions, the flow of blood to a region will 
often be disrupted. The resulting inadequate flow of blood to this region 
will be described in general terms as an insufficiency. The specific type of 
insufficiency is named for the valve involved: aortic insufficiency, mitral 
insufficiency, tricuspid insufficiency, or pulmonary insufficiency. 

If one of the cusps of the valve is forced backward by the force of the 
blood, the condition is referred to as a prolapsed valve. Prolapse may occur 
if the chordae tendineae are damaged or broken, causing the closure 
mechanism to fail. The failure of the valve to close properly disrupts the 
normal one-way flow of blood and results in regurgitation, when the blood 
flows backward from its normal path. Using a stethoscope, the disruption 
to the normal flow of blood produces a heart murmur. 

Stenosis is a condition in which the heart valves become rigid and may 
calcify over time. The loss of flexibility of the valve interferes with normal 
function and may cause the heart to work harder to propel blood through 
the valve, which eventually weakens the heart. Aortic stenosis affects 
approximately 2 percent of the population over 65 years of age, and the 
percentage increases to approximately 4 percent in individuals over 85 
years. Occasionally, one or more of the chordae tendineae will tear or the 
papillary muscle itself may die as a component of a myocardial infarction 
(heart attack). In this case, the patient’s condition will deteriorate 


dramatically and rapidly, and immediate surgical intervention may be 
required. 

Auscultation, or listening to a patient’s heart sounds, is one of the most 
useful diagnostic tools, since it is proven, safe, and inexpensive. The term 
auscultation is derived from the Latin for “to listen,” and the technique has 
been used for diagnostic purposes as far back as the ancient Egyptians. 
Valve and septal disorders will trigger abnormal heart sounds. If a valvular 
disorder is detected or suspected, a test called an echocardiogram, or 
simply an “echo,” may be ordered. Echocardiograms are sonograms of the 
heart and can help in the diagnosis of valve disorders as well as a wide 
variety of heart pathologies. 


Note: 


[ml aC 


Visit this site for a free download, including excellent animations and 
audio of heart sounds. 


Note: 

Career Connection 

Cardiologist 

Cardiologists are medical doctors that specialize in the diagnosis and 
treatment of diseases of the heart. After completing 4 years of medical 
school, cardiologists complete a three-year residency in internal medicine 
followed by an additional three or more years in cardiology. Following this 
10-year period of medical training and clinical experience, they qualify for 
a rigorous two-day examination administered by the Board of Internal 
Medicine that tests their academic training and clinical abilities, including 


diagnostics and treatment. After successful completion of this examination, 
a physician becomes a board-certified cardiologist. Some board-certified 
cardiologists may be invited to become a Fellow of the American College 
of Cardiology (FACC). This professional recognition is awarded to 
outstanding physicians based upon merit, including outstanding 
credentials, achievements, and community contributions to cardiovascular 
medicine. 


Note: 


— 

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Visit this site to learn more about cardiologists. 


Note: 

Career Connection 

Cardiovascular Technologist/Technician 

Cardiovascular technologists/technicians are trained professionals who 
perform a variety of imaging techniques, such as sonograms or 
echocardiograms, used by physicians to diagnose and treat diseases of the 
heart. Nearly all of these positions require an associate degree, and these 
technicians earn a median salary of $49,410 as of May 2010, according to 
the U.S. Bureau of Labor Statistics. Growth within the field is fast, 
projected at 29 percent from 2010 to 2020. 

There is a considerable overlap and complementary skills between cardiac 
technicians and vascular technicians, and so the term cardiovascular 
technician is often used. Special certifications within the field require 
documenting appropriate experience and completing additional and often 
expensive certification examinations. These subspecialties include 


Certified Rhythm Analysis Technician (CRAT), Certified Cardiographic 
Technician (CCT), Registered Congenital Cardiac Sonographer (RCCS), 
Registered Cardiac Electrophysiology Specialist (RCES), Registered 
Cardiovascular Invasive Specialist (RCIS), Registered Cardiac 
Sonographer (RCS), Registered Vascular Specialist (RVS), and Registered 
Phlebology Sonographer (RPhS). 


Note: 
[= [=] 
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a Openstax COLLEGE 
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eo 


Visit this site for more information on cardiovascular 
technologists/technicians. 


Coronary Circulation 


You will recall that the heart is a remarkable pump composed largely of 
cardiac muscle cells that are incredibly active throughout life. Like all other 
cells, a cardiomyocyte requires a reliable supply of oxygen and nutrients, 
and a way to remove wastes, so it needs a dedicated, complex, and 
extensive coronary circulation. And because of the critical and nearly 
ceaseless activity of the heart throughout life, this need for a blood supply is 
even greater than for a typical cell. However, coronary circulation is not 
continuous; rather, it cycles, reaching a peak when the heart muscle is 
relaxed and nearly ceasing while it is contracting. 


Coronary Arteries 


Coronary arteries supply blood to the myocardium and other components 
of the heart. The first portion of the aorta after it arises from the left 
ventricle gives rise to the coronary arteries. There are three dilations in the 
wall of the aorta just superior to the aortic semilunar valve. Two of these, 
the left posterior aortic sinus and anterior aortic sinus, give rise to the left 
and right coronary arteries, respectively. The third sinus, the right posterior 
aortic sinus, typically does not give rise to a vessel. Coronary vessel 
branches that remain on the surface of the artery and follow the sulci are 
called epicardial coronary arteries. 


The left coronary artery distributes blood to the left side of the heart, the left 
atrium and ventricle, and the interventricular septum. The circumflex 
artery arises from the left coronary artery and follows the coronary sulcus 
to the left. Eventually, it will fuse with the small branches of the right 
coronary artery. The larger anterior interventricular artery, also known 
as the left anterior descending artery (LAD), is the second major branch 
arising from the left coronary artery. It follows the anterior interventricular 
sulcus around the pulmonary trunk. Along the way it gives rise to numerous 
smaller branches that interconnect with the branches of the posterior 
interventricular artery, forming anastomoses. An anastomosis is an area 
where vessels unite to form interconnections that normally allow blood to 
circulate to a region even if there may be partial blockage in another 
branch. The anastomoses in the heart are very small. Therefore, this ability 
is somewhat restricted in the heart so a coronary artery blockage often 
results in death of the cells (myocardial infarction) supplied by the 
particular vessel. 


The right coronary artery proceeds along the coronary sulcus and distributes 
blood to the right atrium, portions of both ventricles, and the heart 
conduction system. Normally, one or more marginal arteries arise from the 
right coronary artery inferior to the right atrium. The marginal arteries 
supply blood to the superficial portions of the right ventricle. On the 
posterior surface of the heart, the right coronary artery gives rise to the 
posterior interventricular artery, also known as the posterior descending 
artery. It runs along the posterior portion of the interventricular sulcus 
toward the apex of the heart, giving rise to branches that supply the 
interventricular septum and portions of both ventricles. [link] presents 


views of the coronary circulation from both the anterior and posterior 
views. 
Coronary Circulation 


Aortic arch 
Left coronary 


z t 
Ascending aorta ately 


Pulmonary trunk 


Right coronary 


Circumflex artery 
artery 


Right atrium 
, ; Anterior 
Atrial arteries interventricular 
artery 


Anterior 
cardiac veins : 
Great cardiac 


: vein 
Small cardiac 


vein 


Marginal artery . . 
Coronary sinus Small cardiac 


vein 


Anterior view 


Circumflex artery 


Great cardiac 
vein 


Marginal 
artery 


Posterior 
interventricular 
artery 


Posterior 
cardiac 
vein 


Right 
coronary 
artery 


Middle cardiac Marginal 
vein artery 


Posterior view 


The anterior view of the heart shows the prominent coronary 
surface vessels. The posterior view of the heart shows the 
prominent coronary surface vessels. 


Note: 
Diseases of the... 
Heart: Myocardial Infarction 


Myocardial infarction (MI) is the formal term for what is commonly 
referred to as a heart attack. It normally results from a lack of blood flow 
(ischemia) and oxygen (hypoxia) to a region of the heart, resulting in death 
of the cardiac muscle cells. An MI often occurs when a coronary artery is 
blocked by the buildup of atherosclerotic plaque consisting of lipids, 
cholesterol and fatty acids, and white blood cells, primarily macrophages. 
It can also occur when a portion of an unstable atherosclerotic plaque 
travels through the coronary arterial system and lodges in one of the 
smaller vessels. The resulting blockage restricts the flow of blood and 
oxygen to the myocardium and causes death of the tissue. MIs may be 
triggered by excessive exercise, in which the partially occluded artery is no 
longer able to pump sufficient quantities of blood, or severe stress, which 
may induce spasm of the smooth muscle in the walls of the vessel. 

In the case of acute MI, there is often sudden pain beneath the sternum 
(retrosternal pain) called angina pectoris, often radiating down the left arm 
in males but not in female patients. Until this anomaly between the sexes 
was discovered, many female patients suffering MIs were misdiagnosed 
and sent home. In addition, patients typically present with difficulty 
breathing and shortness of breath (dyspnea), irregular heartbeat 
(palpations), nausea and vomiting, sweating (diaphoresis), anxiety, and 
fainting (syncope), although not all of these symptoms may be present. 
Many of the symptoms are shared with other medical conditions, including 
anxiety attacks and simple indigestion, so differential diagnosis is critical. 
It is estimated that between 22 and 64 percent of MIs present without any 
symptoms. 

An MI can be confirmed by examining the patient’s ECG, which 
frequently reveals alterations in the ST and Q components. Some 
classification schemes of MI are referred to as ST-elevated MI (STEMI) 
and non-elevated MI (non-STEMTI). In addition, echocardiography or 
cardiac magnetic resonance imaging may be employed. Common blood 
tests indicating an MI include elevated levels of creatine kinase MB (an 
enzyme that catalyzes the conversion of creatine to phosphocreatine, 
consuming ATP) and cardiac troponin (the regulatory protein for muscle 
contraction), both of which are released by damaged cardiac muscle cells. 
Immediate treatments for MI are essential and include administering 
supplemental oxygen, aspirin that helps to break up clots, and 
nitroglycerine administered sublingually (under the tongue) to facilitate its 


absorption. Despite its unquestioned success in treatments and use since 
the 1880s, the mechanism of nitroglycerine is still incompletely understood 
but is believed to involve the release of nitric oxide, a known vasodilator, 
and endothelium-derived releasing factor, which also relaxes the smooth 
muscle in the tunica media of coronary vessels. Longer-term treatments 
include injections of thrombolytic agents such as streptokinase that 
dissolve the clot, the anticoagulant heparin, balloon angioplasty and stents 
to open blocked vessels, and bypass surgery to allow blood to pass around 
the site of blockage. If the damage is extensive, coronary replacement with 
a donor heart or coronary assist device, a sophisticated mechanical device 
that supplements the pumping activity of the heart, may be employed. 
Despite the attention, development of artificial hearts to augment the 
severely limited supply of heart donors has proven less than satisfactory 
but will likely improve in the future. 

MIs may trigger cardiac arrest, but the two are not synonymous. Important 
risk factors for MI include cardiovascular disease, age, smoking, high 
blood levels of the low-density lipoprotein (LDL, often referred to as “bad” 
cholesterol), low levels of high-density lipoprotein (HDL, or “good” 
cholesterol), hypertension, diabetes mellitus, obesity, lack of physical 
exercise, chronic kidney disease, excessive alcohol consumption, and use 
of illegal drugs. 


Coronary Veins 


Coronary veins drain the heart and generally parallel the large surface 
arteries (see [link]). The great cardiac vein can be seen initially on the 
surface of the heart following the interventricular sulcus, but it eventually 
flows along the coronary sulcus into the coronary sinus on the posterior 
surface. The great cardiac vein initially parallels the anterior 
interventricular artery and drains the areas supplied by this vessel. It 
receives several major branches, including the posterior cardiac vein, the 
middle cardiac vein, and the small cardiac vein. The posterior cardiac vein 
parallels and drains the areas supplied by the marginal artery branch of the 
circumflex artery. The middle cardiac vein parallels and drains the areas 
supplied by the posterior interventricular artery. The small cardiac vein 


parallels the right coronary artery and drains the blood from the posterior 
surfaces of the right atrium and ventricle. The coronary sinus is a large, 
thin-walled vein on the posterior surface of the heart lying within the 
atrioventricular sulcus and emptying directly into the right atrium. The 
anterior cardiac veins parallel the small cardiac arteries and drain the 
anterior surface of the right ventricle. Unlike these other cardiac veins, it 
bypasses the coronary sinus and drains directly into the right atrium. 


Note: 

Diseases of the... 

Heart: Coronary Artery Disease 

Coronary artery disease is the leading cause of death worldwide. It occurs 
when the buildup of plaque—a fatty material including cholesterol, 
connective tissue, white blood cells, and some smooth muscle cells— 
within the walls of the arteries obstructs the flow of blood and decreases 
the flexibility or compliance of the vessels. This condition is called 
atherosclerosis, a hardening of the arteries that involves the accumulation 
of plaque. As the coronary blood vessels become occluded, the flow of 
blood to the tissues will be restricted, a condition called ischemia that 
causes the cells to receive insufficient amounts of oxygen, called hypoxia. 
[link] shows the blockage of coronary arteries highlighted by the injection 
of dye. Some individuals with coronary artery disease report pain radiating 
from the chest called angina pectoris, but others remain asymptomatic. If 
untreated, coronary artery disease can lead to MI or a heart attack. 
Atherosclerotic Coronary Arteries 


ik ol = 


Blockage of common trunk wy 
of left coronary artery 


In this coronary angiogram (X-ray), the dye 
makes visible two occluded coronary 
arteries. Such blockages can lead to 
decreased blood flow (ischemia) and 
insufficient oxygen (hypoxia) delivered to 
the cardiac tissues. If uncorrected, this can 
lead to cardiac muscle death (myocardial 
infarction). 


The disease progresses slowly and often begins in children and can be seen 
as fatty “streaks” in the vessels. It then gradually progresses throughout 
life. Well-documented risk factors include smoking, family history, 
hypertension, obesity, diabetes, high alcohol consumption, lack of exercise, 
stress, and hyperlipidemia or high circulating levels of lipids in the blood. 
Treatments may include medication, changes to diet and exercise, 
angioplasty with a balloon catheter, insertion of a stent, or coronary bypass 
procedure. 


Angioplasty is a procedure in which the occlusion is mechanically widened 
with a balloon. A specialized catheter with an expandable tip is inserted 
into a superficial vessel, normally in the leg, and then directed to the site of 
the occlusion. At this point, the balloon is inflated to compress the plaque 
material and to open the vessel to increase blood flow. Then, the balloon is 
deflated and retracted. A stent consisting of a specialized mesh is typically 
inserted at the site of occlusion to reinforce the weakened and damaged 
walls. Stent insertions have been routine in cardiology for more than 40 
years. 

Coronary bypass surgery may also be performed. This surgical procedure 
grafts a replacement vessel obtained from another, less vital portion of the 
body to bypass the occluded area. This procedure is clearly effective in 
treating patients experiencing a MI, but overall does not increase longevity. 
Nor does it seem advisable in patients with stable although diminished 
cardiac capacity since frequently loss of mental acuity occurs following the 
procedure. Long-term changes to behavior, emphasizing diet and exercise 
plus a medicine regime tailored to lower blood pressure, lower cholesterol 
and lipids, and reduce clotting are equally as effective. 


Chapter Review 


The heart resides within the pericardial sac and is located in the mediastinal 
space within the thoracic cavity. The pericardial sac consists of two fused 
layers: an outer fibrous capsule and an inner parietal pericardium lined with 
a serous membrane. Between the pericardial sac and the heart is the 
pericardial cavity, which is filled with lubricating serous fluid. The walls of 
the heart are composed of an outer epicardium, a thick myocardium, and an 
inner lining layer of endocardium. The human heart consists of a pair of 
atria, which receive blood and pump it into a pair of ventricles, which pump 
blood into the vessels. The right atrium receives systemic blood relatively 
low in oxygen and pumps it into the right ventricle, which pumps it into the 
pulmonary circuit. Exchange of oxygen and carbon dioxide occurs in the 
lungs, and blood high in oxygen returns to the left atrium, which pumps 
blood into the left ventricle, which in turn pumps blood into the aorta and 
the remainder of the systemic circuit. The septa are the partitions that 


separate the chambers of the heart. They include the interatrial septum, the 
interventricular septum, and the atrioventricular septum. Two of these 
openings are guarded by the atrioventricular valves, the right tricuspid valve 
and the left mitral valve, which prevent the backflow of blood. Each is 
attached to chordae tendineae that extend to the papillary muscles, which 
are extensions of the myocardium, to prevent the valves from being blown 
back into the atria. The pulmonary valve is located at the base of the 
pulmonary trunk, and the left semilunar valve is located at the base of the 
aorta. The right and left coronary arteries are the first to branch off the aorta 
and arise from two of the three sinuses located near the base of the aorta 
and are generally located in the sulci. Cardiac veins parallel the small 
cardiac arteries and generally drain into the coronary sinus. 


Interactive Link Questions 


Exercise: 


Problem: 


Visit this site to observe an echocardiogram of actual heart valves 
opening and closing. Although much of the heart has been “removed” 
from this gif loop so the chordae tendineae are not visible, why is their 
presence more critical for the atrioventricular valves (tricuspid and 
mitral) than the semilunar (aortic and pulmonary) valves? 


Solution: 


The pressure gradient between the atria and the ventricles is much 
greater than that between the ventricles and the pulmonary trunk and 
aorta. Without the presence of the chordae tendineae and papillary 
muscles, the valves would be blown back (prolapsed) into the atria and 
blood would regurgitate. 


Review Questions 


Exercise: 


Problem: 


Which of the following is not important in preventing backflow of 
blood? 


a. chordae tendineae 
b. papillary muscles 
c. AV valves 

d. endocardium 


Solution: 


D 


Exercise: 


Problem: Which valve separates the left atrium from the left ventricle? 


a. mitral 

b. tricuspid 
c. pulmonary 
d. aortic 


Solution: 


A 
Exercise: 
Problem: 


Which of the following lists the valves in the order through which the 
blood flows from the vena cava through the heart? 


a. tricuspid, pulmonary semilunar, bicuspid, aortic semilunar 
b. mitral, pulmonary semilunar, bicuspid, aortic semilunar 
c. aortic semilunar, pulmonary semilunar, tricuspid, bicuspid 


d. bicuspid, aortic semilunar, tricuspid, pulmonary semilunar 


Solution: 


A 
Exercise: 


Problem: 
Which chamber initially receives blood from the systemic circuit? 


a. left atrium 

b. left ventricle 
c. right atrium 
d. right ventricle 


Solution: 


CG 
Exercise: 
Problem: 
The layer secretes chemicals that help to regulate ionic 


environments and strength of contraction and serve as powerful 
vasoconstrictors. 


a. pericardial sac 
b. endocardium 
c. myocardium 
d. epicardium 


Solution: 


B 


Exercise: 


Problem:The myocardium would be the thickest in the 


a. left atrium 

b. left ventricle 
c. right atrium 
d. right ventricle 


Solution: 
B 
Exercise: 
Problem:In which septum is it normal to find openings in the adult? 
a. interatrial septum 
b. interventricular septum 


c. atrioventricular septum 
d. all of the above 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: 


Describe how the valves keep the blood moving in one direction. 


Solution: 


When the ventricles contract and pressure begins to rise in the 
ventricles, there is an initial tendency for blood to flow back 
(regurgitate) to the atria. However, the papillary muscles also contract, 
placing tension on the chordae tendineae and holding the 
atrioventricular valves (tricuspid and mitral) in place to prevent the 
valves from prolapsing and being forced back into the atria. The 
semilunar valves (pulmonary and aortic) lack chordae tendineae and 
papillary muscles, but do not face the same pressure gradients as do 
the atrioventricular valves. As the ventricles relax and pressure drops 
within the ventricles, there is a tendency for the blood to flow 
backward. However, the valves, consisting of reinforced endothelium 
and connective tissue, fill with blood and seal off the opening 
preventing the return of blood. 


Exercise: 
Problem: 


Why is the pressure in the pulmonary circulation lower than in the 
systemic circulation? 


Solution: 


The pulmonary circuit consists of blood flowing to and from the lungs, 
whereas the systemic circuit carries blood to and from the entire body. 
The systemic circuit is far more extensive, consisting of far more 
vessels and offers much greater resistance to the flow of blood, so the 
heart must generate a higher pressure to overcome this resistance. This 
can be seen in the thickness of the myocardium in the ventricles. 


Glossary 


anastomosis 
(plural = anastomoses) area where vessels unite to allow blood to 
circulate even if there may be partial blockage in another branch 


anterior cardiac veins 


vessels that parallel the small cardiac arteries and drain the anterior 
surface of the right ventricle; bypass the coronary sinus and drain 
directly into the right atrium 


anterior interventricular artery 
(also, left anterior descending artery or LAD) major branch of the left 
coronary artery that follows the anterior interventricular sulcus 


anterior interventricular sulcus 
sulcus located between the left and right ventricles on the anterior 
surface of the heart 


aortic valve 
(also, aortic semilunar valve) valve located at the base of the aorta 


atrioventricular septum 
cardiac septum located between the atria and ventricles; 
atrioventricular valves are located here 


atrioventricular valves 
one-way valves located between the atria and ventricles; the valve on 
the right is called the tricuspid valve, and the one on the left is the 
mitral or bicuspid valve 


atrium 
(plural = atria) upper or receiving chamber of the heart that pumps 
blood into the lower chambers just prior to their contraction; the right 
atrium receives blood from the systemic circuit that flows into the right 
ventricle; the left atrium receives blood from the pulmonary circuit 
that flows into the left ventricle 


auricle 
extension of an atrium visible on the superior surface of the heart 


bicuspid valve 
(also, mitral valve or left atrioventricular valve) valve located between 
the left atrium and ventricle; consists of two flaps of tissue 


cardiac notch 
depression in the medial surface of the inferior lobe of the left lung 
where the apex of the heart is located 


cardiac skeleton 
(also, skeleton of the heart) reinforced connective tissue located within 
the atrioventricular septum; includes four rings that surround the 
openings between the atria and ventricles, and the openings to the 
pulmonary trunk and aorta; the point of attachment for the heart valves 


cardiomyocyte 
muscle cell of the heart 


chordae tendineae 
string-like extensions of tough connective tissue that extend from the 
flaps of the atrioventricular valves to the papillary muscles 


circumflex artery 
branch of the left coronary artery that follows coronary sulcus 


coronary arteries 
branches of the ascending aorta that supply blood to the heart; the left 
coronary artery feeds the left side of the heart, the left atrium and 
ventricle, and the interventricular septum; the right coronary artery 
feeds the right atrium, portions of both ventricles, and the heart 
conduction system 


coronary sinus 
large, thin-walled vein on the posterior surface of the heart that lies 
within the atrioventricular sulcus and drains the heart myocardium 
directly into the right atrium 


coronary sulcus 
sulcus that marks the boundary between the atria and ventricles 


coronary veins 
vessels that drain the heart and generally parallel the large surface 
arteries 


endocardium 
innermost layer of the heart lining the heart chambers and heart valves; 
composed of endothelium reinforced with a thin layer of connective 
tissue that binds to the myocardium 


endothelium 
layer of smooth, simple squamous epithelium that lines the 
endocardium and blood vessels 


epicardial coronary arteries 
surface arteries of the heart that generally follow the sulci 


epicardium 
innermost layer of the serous pericardium and the outermost layer of 
the heart wall 


foramen ovale 
opening in the fetal heart that allows blood to flow directly from the 
right atrium to the left atrium, bypassing the fetal pulmonary circuit 


fossa ovalis 
oval-shaped depression in the interatrial septum that marks the former 
location of the foramen ovale 


great cardiac vein 
vessel that follows the interventricular sulcus on the anterior surface of 
the heart and flows along the coronary sulcus into the coronary sinus 
on the posterior surface; parallels the anterior interventricular artery 
and drains the areas supplied by this vessel 


hypertrophic cardiomyopathy 
pathological enlargement of the heart, generally for no known reason 


inferior vena cava 
large systemic vein that returns blood to the heart from the inferior 
portion of the body 


interatrial septum 


cardiac septum located between the two atria; contains the fossa ovalis 
after birth 


interventricular septum 
cardiac septum located between the two ventricles 


left atrioventricular valve 
(also, mitral valve or bicuspid valve) valve located between the left 
atrium and ventricle; consists of two flaps of tissue 


marginal arteries 
branches of the right coronary artery that supply blood to the 
superficial portions of the right ventricle 


mesothelium 
simple squamous epithelial portion of serous membranes, such as the 
superficial portion of the epicardium (the visceral pericardium) and the 
deepest portion of the pericardium (the parietal pericardium) 


middle cardiac vein 
vessel that parallels and drains the areas supplied by the posterior 
interventricular artery; drains into the great cardiac vein 


mitral valve 
(also, left atrioventricular valve or bicuspid valve) valve located 
between the left atrium and ventricle; consists of two flaps of tissue 


moderator band 
band of myocardium covered by endocardium that arises from the 
inferior portion of the interventricular septum in the right ventricle and 
crosses to the anterior papillary muscle; contains conductile fibers that 
carry electrical signals followed by contraction of the heart 


myocardium 
thickest layer of the heart composed of cardiac muscle cells built upon 
a framework of primarily collagenous fibers and blood vessels that 
supply it and the nervous fibers that help to regulate it 


papillary muscle 
extension of the myocardium in the ventricles to which the chordae 
tendineae attach 


pectinate muscles 
muscular ridges seen on the anterior surface of the right atrium 


pericardial cavity 
cavity surrounding the heart filled with a lubricating serous fluid that 
reduces friction as the heart contracts 


pericardial sac 
(also, pericardium) membrane that separates the heart from other 
mediastinal structures; consists of two distinct, fused sublayers: the 
fibrous pericardium and the parietal pericardium 


pericardium 
(also, pericardial sac) membrane that separates the heart from other 
mediastinal structures; consists of two distinct, fused sublayers: the 
fibrous pericardium and the parietal pericardium 


posterior cardiac vein 
vessel that parallels and drains the areas supplied by the marginal 
artery branch of the circumflex artery; drains into the great cardiac 
vein 


posterior interventricular artery 
(also, posterior descending artery) branch of the right coronary artery 
that runs along the posterior portion of the interventricular sulcus 
toward the apex of the heart and gives rise to branches that supply the 
interventricular septum and portions of both ventricles 


posterior interventricular sulcus 
sulcus located between the left and right ventricles on the anterior 
surface of the heart 


pulmonary arteries 


left and right branches of the pulmonary trunk that carry deoxygenated 
blood from the heart to each of the lungs 


pulmonary capillaries 
capillaries surrounding the alveoli of the lungs where gas exchange 
occurs: carbon dioxide exits the blood and oxygen enters 


pulmonary circuit 
blood flow to and from the lungs 


pulmonary trunk 
large arterial vessel that carries blood ejected from the right ventricle; 
divides into the left and right pulmonary arteries 


pulmonary valve 
(also, pulmonary semilunar valve, the pulmonic valve, or the right 
semilunar valve) valve at the base of the pulmonary trunk that prevents 
backflow of blood into the right ventricle; consists of three flaps 


pulmonary veins 
veins that carry highly oxygenated blood into the left atrium, which 
pumps the blood into the left ventricle, which in turn pumps 
oxygenated blood into the aorta and to the many branches of the 
systemic circuit 


right atrioventricular valve 
(also, tricuspid valve) valve located between the right atrium and 
ventricle; consists of three flaps of tissue 


semilunar valves 
valves located at the base of the pulmonary trunk and at the base of the 
aorta 


septum 
(plural = septa) walls or partitions that divide the heart into chambers 


septum primum 


flap of tissue in the fetus that covers the foramen ovale within a few 
seconds after birth 


small cardiac vein 
parallels the right coronary artery and drains blood from the posterior 
surfaces of the right atrium and ventricle; drains into the great cardiac 
vein 


sulcus 
(plural = sulci) fat-filled groove visible on the surface of the heart; 
coronary vessels are also located in these areas 


superior vena cava 
large systemic vein that returns blood to the heart from the superior 
portion of the body 


systemic circuit 
blood flow to and from virtually all of the tissues of the body 


trabeculae carneae 
ridges of muscle covered by endocardium located in the ventricles 


tricuspid valve 
term used most often in clinical settings for the right atrioventricular 
valve 


valve 
in the cardiovascular system, a specialized structure located within the 
heart or vessels that ensures one-way flow of blood 


ventricle 
one of the primary pumping chambers of the heart located in the lower 
portion of the heart; the left ventricle is the major pumping chamber on 
the lower left side of the heart that ejects blood into the systemic 
circuit via the aorta and receives blood from the left atrium; the right 
ventricle is the major pumping chamber on the lower right side of the 
heart that ejects blood into the pulmonary circuit via the pulmonary 
trunk and receives blood from the right atrium 


Cardiac Muscle and Electrical Activity 
By the end of this section, you will be able to: 


e Describe the structure of cardiac muscle 

e Identify and describe the components of the conducting system that 
distributes electrical impulses through the heart 

¢ Compare the effect of ion movement on membrane potential of cardiac 
conductive and contractile cells 

e Relate characteristics of an electrocardiogram to events in the cardiac 
cycle 

e Identify blocks that can interrupt the cardiac cycle 


Recall that cardiac muscle shares a few characteristics with both skeletal 
muscle and smooth muscle, but it has some unique properties of its own. 
Not the least of these exceptional properties is its ability to initiate an 
electrical potential at a fixed rate that spreads rapidly from cell to cell to 
trigger the contractile mechanism. This property is known as 
autorhythmicity. Neither smooth nor skeletal muscle can do this. Even 
though cardiac muscle has autorhythmicity, heart rate is modulated by the 
endocrine and nervous systems. 


There are two major types of cardiac muscle cells: myocardial contractile 
cells and myocardial conducting cells. The myocardial contractile cells 
constitute the bulk (99 percent) of the cells in the atria and ventricles. 
Contractile cells conduct impulses and are responsible for contractions that 
pump blood through the body. The myocardial conducting cells (1 percent 
of the cells) form the conduction system of the heart. Except for Purkinje 
cells, they are generally much smaller than the contractile cells and have 
few of the myofibrils or filaments needed for contraction. Their function is 
similar in many respects to neurons, although they are specialized muscle 
cells. Myocardial conduction cells initiate and propagate the action 
potential (the electrical impulse) that travels throughout the heart and 
triggers the contractions that propel the blood. 


Structure of Cardiac Muscle 


Compared to the giant cylinders of skeletal muscle, cardiac muscle cells, or 
cardiomyocytes, are considerably shorter with much smaller diameters. 
Cardiac muscle also demonstrates striations, the alternating pattern of dark 
A bands and light I bands attributed to the precise arrangement of the 
myofilaments and fibrils that are organized in sarcomeres along the length 
of the cell ({link]a). These contractile elements are virtually identical to 
skeletal muscle. T (transverse) tubules penetrate from the surface plasma 
membrane, the sarcolemma, to the interior of the cell, allowing the 
electrical impulse to reach the interior. The T tubules are only found at the 
Z, discs, whereas in skeletal muscle, they are found at the junction of the A 
and I bands. Therefore, there are one-half as many T tubules in cardiac 
muscle as in skeletal muscle. In addition, the sarcoplasmic reticulum stores 
few calcium ions, so most of the calcium ions must come from outside the 
cells. The result is a slower onset of contraction. Mitochondria are plentiful, 
providing energy for the contractions of the heart. Typically, 
cardiomyocytes have a single, central nucleus, but two or more nuclei may 
be found in some cells. 


Cardiac muscle cells branch freely. A junction between two adjoining cells 
is marked by a critical structure called an intercalated disc, which helps 
support the synchronized contraction of the muscle ([link]b). The 
sarcolemmas from adjacent cells bind together at the intercalated discs. 
They consist of desmosomes, specialized linking proteoglycans, tight 
junctions, and large numbers of gap junctions that allow the passage of ions 
between the cells and help to synchronize the contraction ([link]c). 
Intercellular connective tissue also helps to bind the cells together. The 
importance of strongly binding these cells together is necessitated by the 
forces exerted by contraction. 

Cardiac Muscle 


Intercalated discs 


Intercalated discs 


Mitochondria Intercalated discs 


Gap junction 


Cardiac 
muscle fiber 


(a) 


Desmosome 


— 
A band | band 
(c) 


(a) Cardiac muscle cells have myofibrils composed of 
myofilaments arranged in sarcomeres, T tubules to transmit 
the impulse from the sarcolemma to the interior of the cell, 

numerous mitochondria for energy, and intercalated discs 
that are found at the junction of different cardiac muscle 
cells. (b) A photomicrograph of cardiac muscle cells shows 
the nuclei and intercalated discs. (c) An intercalated disc 
connects cardiac muscle cells and consists of desmosomes 
and gap junctions. LM x 1600. (Micrograph provided by 
the Regents of the University of Michigan Medical School 
© 2012) 


Cardiac muscle undergoes aerobic respiration patterns, primarily 
metabolizing lipids and carbohydrates. Myoglobin, lipids, and glycogen are 
all stored within the cytoplasm. Cardiac muscle cells undergo twitch-type 
contractions with long refractory periods followed by brief relaxation 
periods. The relaxation is essential so the heart can fill with blood for the 
next cycle. The refractory period is very long to prevent the possibility of 
tetany, a condition in which muscle remains involuntarily contracted. In the 
heart, tetany is not compatible with life, since it would prevent the heart 
from pumping blood. 


Note: 

Everyday Connection 

Repair and Replacement 

Damaged cardiac muscle cells have extremely limited abilities to repair 
themselves or to replace dead cells via mitosis. Recent evidence indicates 
that at least some stem cells remain within the heart that continue to divide 
and at least potentially replace these dead cells. However, newly formed or 
repaired cells are rarely as functional as the original cells, and cardiac 
function is reduced. In the event of a heart attack or MI, dead cells are 
often replaced by patches of scar tissue. Autopsies performed on 
individuals who had successfully received heart transplants show some 
proliferation of original cells. If researchers can unlock the mechanism that 
generates new cells and restore full mitotic capabilities to heart muscle, the 
prognosis for heart attack survivors will be greatly enhanced. To date, 
myocardial cells produced within the patient (in situ) by cardiac stem cells 
seem to be nonfunctional, although those grown in Petri dishes (in vitro) do 
beat. Perhaps soon this mystery will be solved, and new advances in 
treatment will be commonplace. 


Conduction System of the Heart 


If embryonic heart cells are separated into a Petri dish and kept alive, each 
is capable of generating its own electrical impulse followed by contraction. 
When two independently beating embryonic cardiac muscle cells are placed 


together, the cell with the higher inherent rate sets the pace, and the impulse 
spreads from the faster to the slower cell to trigger a contraction. As more 
cells are joined together, the fastest cell continues to assume control of the 
rate. A fully developed adult heart maintains the capability of generating its 
own electrical impulse, triggered by the fastest cells, as part of the cardiac 
conduction system. The components of the cardiac conduction system 
include the sinoatrial node, the atrioventricular node, the atrioventricular 
bundle, the atrioventricular bundle branches, and the Purkinje cells ([link]). 
Conduction System of the Heart 


——™ 
Yd ~ 


~. 


Frontal plane = 2 


| 
through heart | Arch of aorta 


= ~~ 


(SS ———— Bachman’s bundle 
<= 


Sinoatrial LL Pe | 
(SA) node 
Anterior internodal 
Atrioventricular 
(AV) node 

Middle internodal 


Posterior internodal 


Left atrium 


Atrioventricular (AV) 
bundle (bundle of His) 


Left ventricle 


Right and left bundle 
branches 


Right atrium 


Right ventricle 


Purkinje fibers 


Anterior view of frontal section 


Specialized conducting components of the heart include 
the sinoatrial node, the internodal pathways, the 
atrioventricular node, the atrioventricular bundle, the 
right and left bundle branches, and the Purkinje fibers. 


Sinoatrial (SA) Node 


Normal cardiac rhythm is established by the sinoatrial (SA) node, a 
specialized clump of myocardial conducting cells located in the superior 


and posterior walls of the right atrium in close proximity to the orifice of 
the superior vena cava. The SA node has the highest inherent rate of 
depolarization and is known as the pacemaker of the heart. It initiates the 
sinus rhythm, or normal electrical pattern followed by contraction of the 
heart. 


This impulse spreads from its initiation in the SA node throughout the atria 
through specialized internodal pathways, to the atrial myocardial 
contractile cells and the atrioventricular node. The internodal pathways 
consist of three bands (anterior, middle, and posterior) that lead directly 
from the SA node to the next node in the conduction system, the 
atrioventricular node (see [link]). The impulse takes approximately 50 ms 
(milliseconds) to travel between these two nodes. The relative importance 
of this pathway has been debated since the impulse would reach the 
atrioventricular node simply following the cell-by-cell pathway through the 
contractile cells of the myocardium in the atria. In addition, there is a 
specialized pathway called Bachmann’s bundle or the interatrial band 
that conducts the impulse directly from the right atrium to the left atrium. 
Regardless of the pathway, as the impulse reaches the atrioventricular 
septum, the connective tissue of the cardiac skeleton prevents the impulse 
from spreading into the myocardial cells in the ventricles except at the 
atrioventricular node. [link] illustrates the initiation of the impulse in the 
SA node that then spreads the impulse throughout the atria to the 
atrioventricular node. 

Cardiac Conduction 


ACS 
=o BN ae 
“aN 
(. |. 


© \ 
7 


(1) The sinoatrial (SA) node and the 
remainder of the conduction system are at 
rest. (2) The SA node initiates the action 
potential, which sweeps across the atria. (3) 
After reaching the atrioventricular node, 
there is a delay of approximately 100 ms 
that allows the atria to complete pumping 
blood before the impulse is transmitted to 
the atrioventricular bundle. (4) Following 
the delay, the impulse travels through the 
atrioventricular bundle and bundle branches 
to the Purkinje fibers, and also reaches the 
right papillary muscle via the moderator 
band. (5) The impulse spreads to the 
contractile fibers of the ventricle. (6) 
Ventricular contraction begins. 


The electrical event, the wave of depolarization, is the trigger for muscular 
contraction. The wave of depolarization begins in the right atrium, and the 
impulse spreads across the superior portions of both atria and then down 
through the contractile cells. The contractile cells then begin contraction 
from the superior to the inferior portions of the atria, efficiently pumping 
blood into the ventricles. 


Atrioventricular (AV) Node 


The atrioventricular (AV) node is a second clump of specialized 
myocardial conductive cells, located in the inferior portion of the right 
atrium within the atrioventricular septum. The septum prevents the impulse 
from spreading directly to the ventricles without passing through the AV 
node. There is a critical pause before the AV node depolarizes and transmits 
the impulse to the atrioventricular bundle (see [link], step 3). This delay in 
transmission is partially attributable to the small diameter of the cells of the 
node, which slow the impulse. Also, conduction between nodal cells is less 
efficient than between conducting cells. These factors mean that it takes the 
impulse approximately 100 ms to pass through the node. This pause is 
critical to heart function, as it allows the atrial cardiomyocytes to complete 
their contraction that pumps blood into the ventricles before the impulse is 
transmitted to the cells of the ventricle itself. With extreme stimulation by 
the SA node, the AV node can transmit impulses maximally at 220 per 
minute. This establishes the typical maximum heart rate in a healthy young 
individual. Damaged hearts or those stimulated by drugs can contract at 
higher rates, but at these rates, the heart can no longer effectively pump 
blood. 


Atrioventricular Bundle (Bundle of His), Bundle Branches, and 
Purkinje Fibers 


Arising from the AV node, the atrioventricular bundle, or bundle of His, 
proceeds through the interventricular septum before dividing into two 
atrioventricular bundle branches, commonly called the left and right 


bundle branches. The left bundle branch has two fascicles. The left bundle 
branch supplies the left ventricle, and the right bundle branch the right 
ventricle. Since the left ventricle is much larger than the right, the left 
bundle branch is also considerably larger than the right. Portions of the right 
bundle branch are found in the moderator band and supply the right 
papillary muscles. Because of this connection, each papillary muscle 
receives the impulse at approximately the same time, so they begin to 
contract simultaneously just prior to the remainder of the myocardial 
contractile cells of the ventricles. This is believed to allow tension to 
develop on the chordae tendineae prior to right ventricular contraction. 
There is no corresponding moderator band on the left. Both bundle 
branches descend and reach the apex of the heart where they connect with 
the Purkinje fibers (see [link], step 4). This passage takes approximately 25 
ms. 


The Purkinje fibers are additional myocardial conductive fibers that spread 
the impulse to the myocardial contractile cells in the ventricles. They 
extend throughout the myocardium from the apex of the heart toward the 
atrioventricular septum and the base of the heart. The Purkinje fibers have a 
fast inherent conduction rate, and the electrical impulse reaches all of the 
ventricular muscle cells in about 75 ms (see [link], step 5). Since the 
electrical stimulus begins at the apex, the contraction also begins at the apex 
and travels toward the base of the heart, similar to squeezing a tube of 
toothpaste from the bottom. This allows the blood to be pumped out of the 
ventricles and into the aorta and pulmonary trunk. The total time elapsed 
from the initiation of the impulse in the SA node until depolarization of the 
ventricles is approximately 225 ms. 


Membrane Potentials and Ion Movement in Cardiac Conductive Cells 


Action potentials are considerably different between cardiac conductive 
cells and cardiac contractive cells. While Na* and K* play essential roles, 
Ca** is also critical for both types of cells. Unlike skeletal muscles and 
neurons, cardiac conductive cells do not have a stable resting potential. 
Conductive cells contain a series of sodium ion channels that allow a 
normal and slow influx of sodium ions that causes the membrane potential 


to rise slowly from an initial value of -60 mV up to about —40 mV. The 
resulting movement of sodium ions creates spontaneous depolarization 
(or prepotential depolarization). At this point, calcium ion channels open 
and Ca** enters the cell, further depolarizing it at a more rapid rate until it 
reaches a value of approximately +5 mV. At this point, the calcium ion 
channels close and K* channels open, allowing outflux of K* and resulting 
in repolarization. When the membrane potential reaches approximately —60 
mV, the K* channels close and Na™ channels open, and the prepotential 
phase begins again. This phenomenon explains the autorhythmicity 
properties of cardiac muscle ({link]). 

Action Potential at the SA Node 


lies Rapid influx of Ca2* 


ae Outflux of Kt 
Depolarization 


fe) Repolarization 


Slow influx of Nat 
Prepotential 


potential 
my) 40 \ Threshold 


-60 


Membrane _90 


—80 


0.8 1.6 
Time (s) 


The prepotential is due to a slow influx of 
sodium ions until the threshold is reached 
followed by a rapid depolarization and 
repolarization. The prepotential accounts for the 
membrane reaching threshold and initiates the 
spontaneous depolarization and contraction of 
the cell. Note the lack of a resting potential. 


Membrane Potentials and Ion Movement in Cardiac Contractile Cells 


There is a distinctly different electrical pattern involving the contractile 
cells. In this case, there is a rapid depolarization, followed by a plateau 


phase and then repolarization. This phenomenon accounts for the long 
refractory periods required for the cardiac muscle cells to pump blood 
effectively before they are capable of firing for a second time. These 
cardiac myocytes normally do not initiate their own electrical potential but 
rather wait for an impulse to reach them. 


Contractile cells demonstrate a much more stable resting phase than 
conductive cells at approximately —80 mV for cells in the atria and -90 mV 
for cells in the ventricles. Despite this initial difference, the other 
components of their action potentials are virtually identical. In both cases, 
when stimulated by an action potential, voltage-gated channels rapidly 
open, beginning the positive-feedback mechanism of depolarization. This 
rapid influx of positively charged ions raises the membrane potential to 
approximately +30 mV, at which point the sodium channels close. The rapid 
depolarization period typically lasts 3-5 ms. Depolarization is followed by 
the plateau phase, in which membrane potential declines relatively slowly. 
This is due in large part to the opening of the slow Ca** channels, allowing 
Ca** to enter the cell while few K* channels are open, allowing K* to exit 
the cell. The relatively long plateau phase lasts approximately 175 ms. Once 
the membrane potential reaches approximately zero, the Ca** channels 
close and K* channels open, allowing K* to exit the cell. The repolarization 
lasts approximately 75 ms. At this point, membrane potential drops until it 
reaches resting levels once more and the cycle repeats. The entire event 
lasts between 250 and 300 ms ([link]). 


The absolute refractory period for cardiac contractile muscle lasts 
approximately 200 ms, and the relative refractory period lasts 
approximately 50 ms, for a total of 250 ms. This extended period is critical, 
since the heart muscle must contract to pump blood effectively and the 
contraction must follow the electrical events. Without extended refractory 
periods, premature contractions would occur in the heart and would not be 
compatible with life. 

Action Potential in Cardiac Contractile Cells 


Na* channels 
close 


Slow Ca?* channels open 


oe Slow Ca2* channels close 


Repolarization 


The plateau 


Rapid depolarization 


K* channels close 
mV 


Voltage-gated 


Refractory period 
ion channels we 
open Absolute Relative 
Influx of Na* 
Time (ms) 
(a) 
Skeletal muscle Cardiac muscle 


Action potential Action potential 


Contraction Contraction 


Tension Tension 


Time (ms) Time (ms) 


(b) 


(a) Note the long plateau phase due to the influx of 
calcium ions. The extended refractory period allows 
the cell to fully contract before another electrical event 
can occur. (b) The action potential for heart muscle is 
compared to that of skeletal muscle. 


Calcium Ions 


Calcium ions play two critical roles in the physiology of cardiac muscle. 
Their influx through slow calcium channels accounts for the prolonged 
plateau phase and absolute refractory period that enable cardiac muscle to 
function properly. Calcium ions also combine with the regulatory protein 
troponin in the troponin-tropomyosin complex; this complex removes the 
inhibition that prevents the heads of the myosin molecules from forming 
cross bridges with the active sites on actin that provide the power stroke of 
contraction. This mechanism is virtually identical to that of skeletal muscle. 
Approximately 20 percent of the calcium required for contraction is 
supplied by the influx of Ca?* during the plateau phase. The remaining Ca?* 
for contraction is released from storage in the sarcoplasmic reticulum. 


Comparative Rates of Conduction System Firing 


The pattern of prepotential or spontaneous depolarization, followed by 
rapid depolarization and repolarization just described, are seen in the SA 
node and a few other conductive cells in the heart. Since the SA node is the 
pacemaker, it reaches threshold faster than any other component of the 
conduction system. It will initiate the impulses spreading to the other 
conducting cells. The SA node, without nervous or endocrine control, 
would initiate a heart impulse approximately 80—100 times per minute. 
Although each component of the conduction system is capable of 
generating its own impulse, the rate progressively slows as you proceed 
from the SA node to the Purkinje fibers. Without the SA node, the AV node 
would generate a heart rate of 40-60 beats per minute. If the AV node were 
blocked, the atrioventricular bundle would fire at a rate of approximately 
30—40 impulses per minute. The bundle branches would have an inherent 
rate of 20—30 impulses per minute, and the Purkinje fibers would fire at 15— 
20 impulses per minute. While a few exceptionally trained aerobic athletes 
demonstrate resting heart rates in the range of 30—40 beats per minute (the 
lowest recorded figure is 28 beats per minute for Miguel Indurain, a 
cyclist), for most individuals, rates lower than 50 beats per minute would 
indicate a condition called bradycardia. Depending upon the specific 
individual, as rates fall much below this level, the heart would be unable to 
maintain adequate flow of blood to vital tissues, initially resulting in 


decreasing loss of function across the systems, unconsciousness, and 
ultimately death. 


Electrocardiogram 


By careful placement of surface electrodes on the body, it is possible to 
record the complex, compound electrical signal of the heart. This tracing of 
the electrical signal is the electrocardiogram (ECG), also commonly 
abbreviated EKG (K coming kardiology, from the German term for 
cardiology). Careful analysis of the ECG reveals a detailed picture of both 
normal and abnormal heart function, and is an indispensable clinical 
diagnostic tool. The standard electrocardiograph (the instrument that 
generates an ECG) uses 3, 5, or 12 leads. The greater the number of leads 
an electrocardiograph uses, the more information the ECG provides. The 
term “lead” may be used to refer to the cable from the electrode to the 
electrical recorder, but it typically describes the voltage difference between 
two of the electrodes. The 12-lead electrocardiograph uses 10 electrodes 
placed in standard locations on the patient’s skin ([link]). In continuous 
ambulatory electrocardiographs, the patient wears a small, portable, battery- 
operated device known as a Holter monitor, or simply a Holter, that 
continuously monitors heart electrical activity, typically for a period of 24 
hours during the patient’s normal routine. 

Standard Placement of ECG Leads 


In a 12-lead ECG, six electrodes are 
placed on the chest, and four 
electrodes are placed on the limbs. 


A normal ECG tracing is presented in [link]. Each component, segment, 
and interval is labeled and corresponds to important electrical events, 
demonstrating the relationship between these events and contraction in the 
heart. 


There are five prominent points on the ECG: the P wave, the QRS complex, 
and the T wave. The small P wave represents the depolarization of the atria. 
The atria begin contracting approximately 25 ms after the start of the P 
wave. The large QRS complex represents the depolarization of the 
ventricles, which requires a much stronger electrical signal because of the 
larger size of the ventricular cardiac muscle. The ventricles begin to 
contract as the QRS reaches the peak of the R wave. Lastly, the T wave 
represents the repolarization of the ventricles. The repolarization of the atria 
occurs during the QRS complex, which masks it on an ECG. 


The major segments and intervals of an ECG tracing are indicated in [link]. 
Segments are defined as the regions between two waves. Intervals include 
one segment plus one or more waves. For example, the PR segment begins 
at the end of the P wave and ends at the beginning of the QRS complex. 
The PR interval starts at the beginning of the P wave and ends with the 
beginning of the QRS complex. The PR interval is more clinically relevant, 
as it measures the duration from the beginning of atrial depolarization (the 
P wave) to the initiation of the QRS complex. Since the Q wave may be 
difficult to view in some tracings, the measurement is often extended to the 
R that is more easily visible. Should there be a delay in passage of the 
impulse from the SA node to the AV node, it would be visible in the PR 
interval. [link] correlates events of heart contraction to the corresponding 
segments and intervals of an ECG. 


Note: 


fe 


— 
meee, <OPENStAX COLLEGE 


Visit this site for a more detailed analysis of ECGs. 


Electrocardiogram 


5mm 


P-R S-T 
segment segment 


Millivolts 


QRS complex 


PR interval QT interval 


A normal tracing shows the P wave, QRS 
complex, and T wave. Also indicated are the 
PR, QT, QRS, and ST intervals, plus the P-R 

and S-T segments. 


ECG Tracing Correlated to the Cardiac Cycle 


0) 


This diagram correlates an ECG tracing with the 
electrical and mechanical events of a heart contraction. 
Each segment of an ECG tracing corresponds to one 
event in the cardiac cycle. 


Note: 

Everyday Connection 

ECG Abnormalities 

Occassionally, an area of the heart other than the SA node will initiate an 
impulse that will be followed by a premature contraction. Such an area, 
which may actually be a component of the conduction system or some 
other contractile cells, is known as an ectopic focus or ectopic pacemaker. 
An ectopic focus may be stimulated by localized ischemia; exposure to 
certain drugs, including caffeine, digitalis, or acetylcholine; elevated 
stimulation by both sympathetic or parasympathetic divisions of the 
autonomic nervous system; or a number of disease or pathological 
conditions. Occasional occurances are generally transitory and nonlife 
threatening, but if the condition becomes chronic, it may lead to either an 
arrhythmia, a deviation from the normal pattern of impulse conduction and 
contraction, or to fibrillation, an uncoordinated beating of the heart. 
While interpretation of an ECG is possible and extremely valuable after 
some training, a full understanding of the complexities and intricacies 
generally requires several years of experience. In general, the size of the 
electrical variations, the duration of the events, and detailed vector analysis 
provide the most comprehensive picture of cardiac function. For example, 
an amplified P wave may indicate enlargement of the atria, an enlarged Q 
wave may indicate a MI, and an enlarged suppressed or inverted Q wave 
often indicates enlarged ventricles. T waves often appear flatter when 
insufficient oxygen is being delivered to the myocardium. An elevation of 
the ST segment above baseline is often seen in patients with an acute MI, 
and may appear depressed below the baseline when hypoxia is occurring. 
As useful as analyzing these electrical recordings may be, there are 
limitations. For example, not all areas suffering a MI may be obvious on 


the ECG. Additionally, it will not reveal the effectiveness of the pumping, 
which requires further testing, such as an ultrasound test called an 
echocardiogram or nuclear medicine imaging. It is also possible for there 
to be pulseless electrical activity, which will show up on an ECG tracing, 
although there is no corresponding pumping action. Common 
abnormalities that may be detected by the ECGs are shown in [link]. 
Common ECG Abnormalities 

Note how half of the P waves 

are not followed by the QRS 

complex and T waves while the 

other half are. 

Question: What would you 


expect to happen to heart rate 
(pulse)? 


| 
| 


(a) Second-degree (partial) block 


| | Note the abnormal electrical 
} | pattern prior to the QRS 

| } complexes. Also note how the 
frequency between the QRS 
complexes has increased. 
~ Question: What would you 
_ expect to happen to heart rate 
(pulse)? 


(b) Atrial fibrillation 


Note the unusual shape of the 
QRS complex, focusing on the 
“S” component. 

Question: What would you 
expect to happen to heart rate 
(pulse)? 


(c 


Ventricular tachycardia 
Note the total lack of normal 


electrical activity. 
Question: What would you 


_ expect to happen to heart 
| | rate (pulse)? 
| } | 


(d) Ventricular fibrillation 


Note that in a third-degree block 
some of the impulses initiated by 
the SA node do not reach the 
AV node while others do. Also note 
that the P waves are not followed 
by the QRS complex. 

| | | / Question: What would you expect 
(e) Third-degree block to happen to heart rate (pulse)? 


(a) In a second-degree or partial block, one-half of the P 
waves are not followed by the QRS complex and T waves 
while the other half are. (b) In atrial fibrillation, the 
electrical pattern is abnormal prior to the QRS complex, 
and the frequency between the QRS complexes has 
increased. (c) In ventricular tachycardia, the shape of the 


QRS complex is abnormal. (d) In ventricular fibrillation, 

there is no normal electrical activity. (e) In a third-degree 

block, there is no correlation between atrial activity (the P 
wave) and ventricular activity (the QRS complex). 


Visit this site for a more complete library of abnormal ECGs. 


Note: 

Everyday Connection 

External Automated Defibrillators 

In the event that the electrical activity of the heart is severely disrupted, 
cessation of electrical activity or fibrillation may occur. In fibrillation, the 
heart beats in a wild, uncontrolled manner, which prevents it from being 
able to pump effectively. Atrial fibrillation (see [link]b) is a serious 
condition, but as long as the ventricles continue to pump blood, the 
patient’s life may not be in immediate danger. Ventricular fibrillation (see 
[link ]d) is a medical emergency that requires life support, because the 
ventricles are not effectively pumping blood. In a hospital setting, it is 
often described as “code blue.” If untreated for as little as a few minutes, 
ventricular fibrillation may lead to brain death. The most common 
treatment is defibrillation, which uses special paddles to apply a charge to 
the heart from an external electrical source in an attempt to establish a 
normal sinus rhythm ([link]). A defibrillator effectively stops the heart so 


that the SA node can trigger a normal conduction cycle. Because of their 
effectiveness in reestablishing a normal sinus rhythm, external automated 
defibrillators (EADs) are being placed in areas frequented by large 
numbers of people, such as schools, restaurants, and airports. These 
devices contain simple and direct verbal instructions that can be followed 
by nonmedical personnel in an attempt to save a life. 

Defibrillators 


N 
\ 


(a) An external automatic defibrillator can be used by 
nonmedical personnel to reestablish a normal sinus rhythm 
in a person with fibrillation. (b) Defibrillator paddles are 
more commonly used in hospital settings. (credit b: 
“widerider107”/flickr.com) 


A heart block refers to an interruption in the normal conduction pathway. 
The nomenclature for these is very straightforward. SA nodal blocks occur 
within the SA node. AV nodal blocks occur within the AV node. Infra- 
Hisian blocks involve the bundle of His. Bundle branch blocks occur within 
either the left or right atrioventricular bundle branches. Hemiblocks are 
partial and occur within one or more fascicles of the atrioventricular bundle 
branch. Clinically, the most common types are the AV nodal and infra- 
Hisian blocks. 


AV blocks are often described by degrees. A first-degree or partial block 
indicates a delay in conduction between the SA and AV nodes. This can be 
recognized on the ECG as an abnormally long PR interval. A second-degree 
or incomplete block occurs when some impulses from the SA node reach 
the AV node and continue, while others do not. In this instance, the ECG 
would reveal some P waves not followed by a QRS complex, while others 
would appear normal. In the third-degree or complete block, there is no 
correlation between atrial activity (the P wave) and ventricular activity (the 
QRS complex). Even in the event of a total SA block, the AV node will 
assume the role of pacemaker and continue initiating contractions at 40-60 
contractions per minute, which is adequate to maintain consciousness. 
Second- and third-degree blocks are demonstrated on the ECG presented in 
[link]. 


When arrhythmias become a chronic problem, the heart maintains a 
junctional rhythm, which originates in the AV node. In order to speed up the 
heart rate and restore full sinus rhythm, a cardiologist can implant an 
artificial pacemaker, which delivers electrical impulses to the heart 
muscle to ensure that the heart continues to contract and pump blood 
effectively. These artificial pacemakers are programmable by the 
cardiologists and can either provide stimulation temporarily upon demand 
or on a continuous basis. Some devices also contain built-in defibrillators. 


Cardiac Muscle Metabolism 


Normally, cardiac muscle metabolism is entirely aerobic. Oxygen from the 
lungs is brought to the heart, and every other organ, attached to the 
hemoglobin molecules within the erythrocytes. Heart cells also store 
appreciable amounts of oxygen in myoglobin. Normally, these two 
mechanisms, circulating oxygen and oxygen attached to myoglobin, can 
supply sufficient oxygen to the heart, even during peak performance. 


Fatty acids and glucose from the circulation are broken down within the 
mitochondria to release energy in the form of ATP. Both fatty acid droplets 
and glycogen are stored within the sarcoplasm and provide additional 
nutrient supply. (Seek additional content for more detail about metabolism.) 


Chapter Review 


The heart is regulated by both neural and endocrine control, yet it is capable 
of initiating its own action potential followed by muscular contraction. The 
conductive cells within the heart establish the heart rate and transmit it 
through the myocardium. The contractile cells contract and propel the 
blood. The normal path of transmission for the conductive cells is the 
sinoatrial (SA) node, internodal pathways, atrioventricular (AV) node, 
atrioventricular (AV) bundle of His, bundle branches, and Purkinje fibers. 
The action potential for the conductive cells consists of a prepotential phase 
with a slow influx of Na* followed by a rapid influx of Ca** and outflux of 
K*. Contractile cells have an action potential with an extended plateau 
phase that results in an extended refractory period to allow complete 
contraction for the heart to pump blood effectively. Recognizable points on 
the ECG include the P wave that corresponds to atrial depolarization, the 
QRS complex that corresponds to ventricular depolarization, and the T 
wave that corresponds to ventricular repolarization. 


Review Questions 


Exercise: 


Problem: Which of the following is unique to cardiac muscle cells? 


a. Only cardiac muscle contains a sarcoplasmic reticulum. 

b. Only cardiac muscle has gap junctions. 

c. Only cardiac muscle is capable of autorhythmicity 

d. Only cardiac muscle has a high concentration of mitochondria. 


Solution: 


c 


Exercise: 


Problem:The influx of which ion accounts for the plateau phase? 


a. sodium 

b. potassium 
c. chloride 
d. calcium 


Solution: 


D 
Exercise: 


Problem: 
Which portion of the ECG corresponds to repolarization of the atria? 


a. P wave 

b. QRS complex 

c. T wave 

d. none of the above: atrial repolarization is masked by ventricular 
depolarization 


Solution: 


D 
Exercise: 
Problem: 


Which component of the heart conduction system would have the 
slowest rate of firing? 


a. atrioventricular node 
b. atrioventricular bundle 
c. bundle branches 

d. Purkinje fibers 


Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: 
Why is the plateau phase so critical to cardiac muscle function? 
Solution: 


It prevents additional impulses from spreading through the heart 
prematurely, thereby allowing the muscle sufficient time to contract 
and pump blood effectively. 


Exercise: 
Problem: 


How does the delay of the impulse at the atrioventricular node 
contribute to cardiac function? 


Solution: 


It ensures sufficient time for the atrial muscle to contract and pump 
blood into the ventricles prior to the impulse being conducted into the 
lower chambers. 


Exercise: 


Problem: 


How do gap junctions and intercalated disks aid contraction of the 
heart? 


Solution: 


Gap junctions within the intercalated disks allow impulses to spread 
from one cardiac muscle cell to another, allowing sodium, potassium, 


and calcium ions to flow between adjacent cells, propagating the 
action potential, and ensuring coordinated contractions. 


Exercise: 


Problem: 


Why do the cardiac muscles cells demonstrate autorhythmicity? 


Solution: 


Without a true resting potential, there is a slow influx of sodium ions 
through slow channels that produces a prepotential that gradually 
reaches threshold. 


Glossary 


artificial pacemaker 
medical device that transmits electrical signals to the heart to ensure 
that it contracts and pumps blood to the body 


atrioventricular bundle 
(also, bundle of His) group of specialized myocardial conductile cells 
that transmit the impulse from the AV node through the 
interventricular septum; form the left and right atrioventricular bundle 
branches 


atrioventricular bundle branches 
(also, left or right bundle branches) specialized myocardial conductile 
cells that arise from the bifurcation of the atrioventricular bundle and 
pass through the interventricular septum; lead to the Purkinje fibers 
and also to the right papillary muscle via the moderator band 


atrioventricular (AV) node 
clump of myocardial cells located in the inferior portion of the right 
atrium within the atrioventricular septum; receives the impulse from 
the SA node, pauses, and then transmits it into specialized conducting 
cells within the interventricular septum 


autorhythmicity 
ability of cardiac muscle to initiate its own electrical impulse that 
triggers the mechanical contraction that pumps blood at a fixed pace 
without nervous or endocrine control 


Bachmann’s bundle 
(also, interatrial band) group of specialized conducting cells that 
transmit the impulse directly from the SA node in the right atrium to 
the left atrium 


bundle of His 
(also, atrioventricular bundle) group of specialized myocardial 
conductile cells that transmit the impulse from the AV node through 
the interventricular septum; form the left and right atrioventricular 
bundle branches 


electrocardiogram (ECG) 
surface recording of the electrical activity of the heart that can be used 
for diagnosis of irregular heart function; also abbreviated as EKG 


heart block 
interruption in the normal conduction pathway 


interatrial band 
(also, Bachmann’s bundle) group of specialized conducting cells that 
transmit the impulse directly from the SA node in the right atrium to 
the left atrium 


intercalated disc 
physical junction between adjacent cardiac muscle cells; consisting of 
desmosomes, specialized linking proteoglycans, and gap junctions that 
allow passage of ions between the two cells 


internodal pathways 
specialized conductile cells within the atria that transmit the impulse 
from the SA node throughout the myocardial cells of the atrium and to 
the AV node 


myocardial conducting cells 
specialized cells that transmit electrical impulses throughout the heart 
and trigger contraction by the myocardial contractile cells 


myocardial contractile cells 
bulk of the cardiac muscle cells in the atria and ventricles that conduct 
impulses and contract to propel blood 


P wave 
component of the electrocardiogram that represents the depolarization 
of the atria 


pacemaker 
cluster of specialized myocardial cells known as the SA node that 
initiates the sinus rhythm 


prepotential depolarization 
(also, spontaneous depolarization) mechanism that accounts for the 
autorhythmic property of cardiac muscle; the membrane potential 
increases as sodium ions diffuse through the always-open sodium ion 
channels and causes the electrical potential to rise 


Purkinje fibers 
specialized myocardial conduction fibers that arise from the bundle 
branches and spread the impulse to the myocardial contraction fibers 
of the ventricles 


QRS complex 
component of the electrocardiogram that represents the depolarization 
of the ventricles and includes, as a component, the repolarization of the 
atria 


sinoatrial (SA) node 
known as the pacemaker, a specialized clump of myocardial 
conducting cells located in the superior portion of the right atrium that 
has the highest inherent rate of depolarization that then spreads 
throughout the heart 


sinus rhythm 
normal contractile pattern of the heart 


spontaneous depolarization 
(also, prepotential depolarization) the mechanism that accounts for the 
autorhythmic property of cardiac muscle; the membrane potential 
increases as sodium ions diffuse through the always-open sodium ion 
channels and causes the electrical potential to rise 


T wave 
component of the electrocardiogram that represents the repolarization 


of the ventricles 


Cardiac Cycle 
By the end of this section, you will be able to: 


e Describe the relationship between blood pressure and blood flow 

e Summarize the events of the cardiac cycle 

e¢ Compare atrial and ventricular systole and diastole 

e Relate heart sounds detected by auscultation to action of heart’s valves 


The period of time that begins with contraction of the atria and ends with 
ventricular relaxation is known as the cardiac cycle ([link]). The period of 
contraction that the heart undergoes while it pumps blood into circulation is 
called systole. The period of relaxation that occurs as the chambers fill with 
blood is called diastole. Both the atria and ventricles undergo systole and 
diastole, and it is essential that these components be carefully regulated and 
coordinated to ensure blood is pumped efficiently to the body. 

Overview of the Cardiac Cycle 


| 


The cardiac cycle begins with atrial systole and 
progresses to ventricular systole, atrial 
diastole, and ventricular diastole, when the 
cycle begins again. Correlations to the ECG 
are highlighted. 


Pressures and Flow 


Fluids, whether gases or liquids, are materials that flow according to 
pressure gradients—that is, they move from regions that are higher in 
pressure to regions that are lower in pressure. Accordingly, when the heart 
chambers are relaxed (diastole), blood will flow into the atria from the 
veins, which are higher in pressure. As blood flows into the atria, the 
pressure will rise, so the blood will initially move passively from the atria 
into the ventricles. When the action potential triggers the muscles in the 
atria to contract (atrial systole), the pressure within the atria rises further, 
pumping blood into the ventricles. During ventricular systole, pressure rises 
in the ventricles, pumping blood into the pulmonary trunk from the right 
ventricle and into the aorta from the left ventricle. Again, as you consider 
this flow and relate it to the conduction pathway, the elegance of the system 
should become apparent. 


Phases of the Cardiac Cycle 


At the beginning of the cardiac cycle, both the atria and ventricles are 
relaxed (diastole). Blood is flowing into the right atrium from the superior 
and inferior venae cavae and the coronary sinus. Blood flows into the left 
atrium from the four pulmonary veins. The two atrioventricular valves, the 
tricuspid and mitral valves, are both open, so blood flows unimpeded from 
the atria and into the ventricles. Approximately 70—80 percent of ventricular 
filling occurs by this method. The two semilunar valves, the pulmonary and 
aortic valves, are closed, preventing backflow of blood into the right and 
left ventricles from the pulmonary trunk on the right and the aorta on the 
left. 


Atrial Systole and Diastole 


Contraction of the atria follows depolarization, represented by the P wave 
of the ECG. As the atrial muscles contract from the superior portion of the 
atria toward the atrioventricular septum, pressure rises within the atria and 
blood is pumped into the ventricles through the open atrioventricular 
(tricuspid, and mitral or bicuspid) valves. At the start of atrial systole, the 
ventricles are normally filled with approximately 70-80 percent of their 
capacity due to inflow during diastole. Atrial contraction, also referred to as 
the “atrial kick,” contributes the remaining 20—30 percent of filling (see 
[link]). Atrial systole lasts approximately 100 ms and ends prior to 
ventricular systole, as the atrial muscle returns to diastole. 


Ventricular Systole 


Ventricular systole (see [link]) follows the depolarization of the ventricles 
and is represented by the QRS complex in the ECG. It may be conveniently 
divided into two phases, lasting a total of 270 ms. At the end of atrial 
systole and just prior to atrial contraction, the ventricles contain 
approximately 130 mL blood in a resting adult in a standing position. This 
volume is known as the end diastolic volume (EDV) or preload. 


Initially, as the muscles in the ventricle contract, the pressure of the blood 
within the chamber rises, but it is not yet high enough to open the semilunar 
(pulmonary and aortic) valves and be ejected from the heart. However, 
blood pressure quickly rises above that of the atria that are now relaxed and 
in diastole. This increase in pressure causes blood to flow back toward the 
atria, closing the tricuspid and mitral valves. Since blood is not being 
ejected from the ventricles at this early stage, the volume of blood within 
the chamber remains constant. Consequently, this initial phase of 
ventricular systole is known as isovolumic contraction, also called 
isovolumetric contraction (see [link]). 


In the second phase of ventricular systole, the ventricular ejection phase, 
the contraction of the ventricular muscle has raised the pressure within the 
ventricle to the point that it is greater than the pressures in the pulmonary 


trunk and the aorta. Blood is pumped from the heart, pushing open the 
pulmonary and aortic semilunar valves. Pressure generated by the left 
ventricle will be appreciably greater than the pressure generated by the right 
ventricle, since the existing pressure in the aorta will be so much higher. 
Nevertheless, both ventricles pump the same amount of blood. This 
quantity is referred to as stroke volume. Stroke volume will normally be in 
the range of 70-80 mL. Since ventricular systole began with an EDV of 
approximately 130 mL of blood, this means that there is still 50-60 mL of 
blood remaining in the ventricle following contraction. This volume of 
blood is known as the end systolic volume (ESV). 


Ventricular Diastole 


Ventricular relaxation, or diastole, follows repolarization of the ventricles 
and is represented by the T wave of the ECG. It too is divided into two 
distinct phases and lasts approximately 430 ms. 


During the early phase of ventricular diastole, as the ventricular muscle 
relaxes, pressure on the remaining blood within the ventricle begins to fall. 
When pressure within the ventricles drops below pressure in both the 
pulmonary trunk and aorta, blood flows back toward the heart, producing 
the dicrotic notch (small dip) seen in blood pressure tracings. The semilunar 
valves close to prevent backflow into the heart. Since the atrioventricular 
valves remain closed at this point, there is no change in the volume of blood 
in the ventricle, so the early phase of ventricular diastole is called the 
isovolumic ventricular relaxation phase, also called isovolumetric 
ventricular relaxation phase (see [link]). 


In the second phase of ventricular diastole, called late ventricular diastole, 
as the ventricular muscle relaxes, pressure on the blood within the 
ventricles drops even further. Eventually, it drops below the pressure in the 
atria. When this occurs, blood flows from the atria into the ventricles, 
pushing open the tricuspid and mitral valves. As pressure drops within the 
ventricles, blood flows from the major veins into the relaxed atria and from 
there into the ventricles. Both chambers are in diastole, the atrioventricular 


valves are open, and the semilunar valves remain closed (see [link]). The 
cardiac cycle is complete. 


[link] illustrates the relationship between the cardiac cycle and the ECG. 
Relationship between the Cardiac Cycle and ECG 
R 


One cardiac cycle 


Initially, both the atria and ventricles are relaxed 
(diastole). The P wave represents depolarization of the 
atria and is followed by atrial contraction (systole). 
Atrial systole extends until the QRS complex, at 
which point, the atria relax. The QRS complex 
represents depolarization of the ventricles and is 
followed by ventricular contraction. The T wave 
represents the repolarization of the ventricles and 
marks the beginning of ventricular relaxation. 


Heart Sounds 


One of the simplest, yet effective, diagnostic techniques applied to assess 
the state of a patient’s heart is auscultation using a stethoscope. 


In a normal, healthy heart, there are only two audible heart sounds: S, and 
S>. S; is the sound created by the closing of the atrioventricular valves 
during ventricular contraction and is normally described as a “lub,” or first 
heart sound. The second heart sound, Sp, is the sound of the closing of the 


semilunar valves during ventricular diastole and is described as a “dub” 
({link]). In both cases, as the valves close, the openings within the 
atrioventricular septum guarded by the valves will become reduced, and 
blood flow through the opening will become more turbulent until the valves 
are fully closed. There is a third heart sound, S3, but it is rarely heard in 
healthy individuals. It may be the sound of blood flowing into the atria, or 
blood sloshing back and forth in the ventricle, or even tensing of the 
chordae tendineae. S3 may be heard in youth, some athletes, and pregnant 
women. If the sound is heard later in life, it may indicate congestive heart 
failure, warranting further tests. Some cardiologists refer to the collective 
S1, Sz, and S3 sounds as the “Kentucky gallop,” because they mimic those 
produced by a galloping horse. The fourth heart sound, Sy, results from the 
contraction of the atria pushing blood into a stiff or hypertrophic ventricle, 
indicating failure of the left ventricle. S, occurs prior to S; and the 
collective sounds S,, S;, and S> are referred to by some cardiologists as the 
“Tennessee gallop,” because of their similarity to the sound produced by a 
galloping horse with a different gait. A few individuals may have both S3 
and Sy, and this combined sound is referred to as Sv. 

Heart Sounds and the Cardiac Cycle 


Semilunar 


120 valves close 


Semilunar 
100 valves open 


aS 


Aortic pressure 


Ventricular pressure 


Pressure (mm Hg) 
(op) 
ro] 


AV valves 
open 


/ 


AV valves 
close 


Atrial pressure 


1st 2nd 3rd 
Heart sounds |), 


“| ub” “Dub” 


In this illustration, the x-axis reflects time with a 


recording of the heart sounds. The y-axis represents 
pressure. 


The term murmur is used to describe an unusual sound coming from the 
heart that is caused by the turbulent flow of blood. Murmurs are graded on a 
scale of 1 to 6, with 1 being the most common, the most difficult sound to 
detect, and the least serious. The most severe is a 6. Phonocardiograms or 
auscultograms can be used to record both normal and abnormal sounds 
using specialized electronic stethoscopes. 


During auscultation, it is common practice for the clinician to ask the 
patient to breathe deeply. This procedure not only allows for listening to 
airflow, but it may also amplify heart murmurs. Inhalation increases blood 
flow into the right side of the heart and may increase the amplitude of right- 
sided heart murmurs. Expiration partially restricts blood flow into the left 
side of the heart and may amplify left-sided heart murmurs. [link] indicates 
proper placement of the bell of the stethoscope to facilitate auscultation. 
Stethoscope Placement for Auscultation 


Aortic valve Pulmonary valve 


Tricuspid valve Mitral valve 


Proper placement of the bell of the stethoscope 
facilitates auscultation. At each of the four locations 
on the chest, a different valve can be heard. 


Chapter Review 


The cardiac cycle comprises a complete relaxation and contraction of both 
the atria and ventricles, and lasts approximately 0.8 seconds. Beginning 
with all chambers in diastole, blood flows passively from the veins into the 
atria and past the atrioventricular valves into the ventricles. The atria begin 
to contract (atrial systole), following depolarization of the atria, and pump 
blood into the ventricles. The ventricles begin to contract (ventricular 
systole), raising pressure within the ventricles. When ventricular pressure 
rises above the pressure in the atria, blood flows toward the atria, producing 
the first heart sound, S, or lub. As pressure in the ventricles rises above two 
major arteries, blood pushes open the two semilunar valves and moves into 
the pulmonary trunk and aorta in the ventricular ejection phase. Following 
ventricular repolarization, the ventricles begin to relax (ventricular 
diastole), and pressure within the ventricles drops. As ventricular pressure 
drops, there is a tendency for blood to flow back into the atria from the 
major arteries, producing the dicrotic notch in the ECG and closing the two 
semilunar valves. The second heart sound, S» or dub, occurs when the 
semilunar valves close. When the pressure falls below that of the atria, 
blood moves from the atria into the ventricles, opening the atrioventricular 
valves and marking one complete heart cycle. The valves prevent backflow 
of blood. Failure of the valves to operate properly produces turbulent blood 
flow within the heart; the resulting heart murmur can often be heard with a 
stethoscope. 


Review Questions 


Exercise: 


Problem: 


The cardiac cycle consists of a distinct relaxation and contraction 
phase. Which term is typically used to refer ventricular contraction 
while no blood is being ejected? 


a. systole 

b. diastole 

c. quiescent 

d. isovolumic contraction 


Solution: 


D 


Exercise: 


Problem: Most blood enters the ventricle during 


a. atrial systole 

b. atrial diastole 

c. ventricular systole 

d. isovolumic contraction 


Solution: 


B 
Exercise: 


Problem: 
The first heart sound represents which portion of the cardiac cycle? 


a. atrial systole 
b. ventricular systole 
c. closing of the atrioventricular valves 


d. closing of the semilunar valves 


Solution: 


‘s 


Exercise: 


Problem: Ventricular relaxation immediately follows 


a. atrial depolarization 
b. ventricular repolarization 
c. ventricular depolarization 
d. atrial repolarization 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


Describe one cardiac cycle, beginning with both atria and ventricles 
relaxed. 


Solution: 


The cardiac cycle comprises a complete relaxation and contraction of 
both the atria and ventricles, and lasts approximately 0.8 seconds. 
Beginning with all chambers in diastole, blood flows passively from 
the veins into the atria and past the atrioventricular valves into the 
ventricles. The atria begin to contract following depolarization of the 
atria and pump blood into the ventricles. The ventricles begin to 


contract, raising pressure within the ventricles. When ventricular 
pressure rises above the pressure in the two major arteries, blood 
pushes open the two semilunar valves and moves into the pulmonary 
trunk and aorta in the ventricular ejection phase. Following ventricular 
repolarization, the ventricles begin to relax, and pressure within the 
ventricles drops. When the pressure falls below that of the atria, blood 
moves from the atria into the ventricles, opening the atrioventricular 
valves and marking one complete heart cycle. 


Glossary 


cardiac cycle 
period of time between the onset of atrial contraction (atrial systole) 
and ventricular relaxation (ventricular diastole) 


diastole 
period of time when the heart muscle is relaxed and the chambers fill 
with blood 


end diastolic volume (EDV) 
(also, preload) the amount of blood in the ventricles at the end of atrial 
systole just prior to ventricular contraction 


end systolic volume (ESV) 
amount of blood remaining in each ventricle following systole 


heart sounds 
sounds heard via auscultation with a stethoscope of the closing of the 
atrioventricular valves (“lub”) and semilunar valves (“dub”’) 


isovolumic contraction 
(also, isovolumetric contraction) initial phase of ventricular 
contraction in which tension and pressure in the ventricle increase, but 
no blood is pumped or ejected from the heart 


isovolumic ventricular relaxation phase 


initial phase of the ventricular diastole when pressure in the ventricles 
drops below pressure in the two major arteries, the pulmonary trunk, 
and the aorta, and blood attempts to flow back into the ventricles, 
producing the dicrotic notch of the ECG and closing the two semilunar 
valves 


murmur 
unusual heart sound detected by auscultation; typically related to septal 
or valve defects 


preload 
(also, end diastolic volume) amount of blood in the ventricles at the 
end of atrial systole just prior to ventricular contraction 


systole 
period of time when the heart muscle is contracting 


ventricular ejection phase 
second phase of ventricular systole during which blood is pumped 
from the ventricle 


Structure and Function of Blood Vessels 
By the end of this section, you will be able to: 


e Compare and contrast the three tunics that make up the walls of most 
blood vessels 

e Distinguish between elastic arteries, muscular arteries, and arterioles 
on the basis of structure, location, and function 

e Describe the basic structure of a capillary bed, from the supplying 
metarteriole to the venule into which it drains 

e Explain the structure and function of venous valves in the large veins 
of the extremities 


Blood is carried through the body via blood vessels. An artery is a blood 
vessel that carries blood away from the heart, where it branches into ever- 
smaller vessels. Eventually, the smallest arteries, vessels called arterioles, 
further branch into tiny capillaries, where nutrients and wastes are 
exchanged, and then combine with other vessels that exit capillaries to form 
venules, small blood vessels that carry blood to a vein, a larger blood vessel 
that returns blood to the heart. 


Arteries and veins transport blood in two distinct circuits: the systemic 
circuit and the pulmonary circuit ([link]). Systemic arteries provide blood 
rich in oxygen to the body’s tissues. The blood returned to the heart through 
systemic veins has less oxygen, since much of the oxygen carried by the 
arteries has been delivered to the cells. In contrast, in the pulmonary circuit, 
arteries carry blood low in oxygen exclusively to the lungs for gas 
exchange. Pulmonary veins then return freshly oxygenated blood from the 
lungs to the heart to be pumped back out into systemic circulation. 
Although arteries and veins differ structurally and functionally, they share 
certain features. 

Cardiovascular Circulation 


Lungs 


2c 
GO : 
5 & Pulmonary Pulmonary vein 
=) 
§3 artery 
as 
Vena cava Aorta 
Upper body 
Liver 
Hepatic vein Hepatic artery 


Hepatic portal vein 


Systemic 
circulation 


Stomach, 
intestines Hi Vessels transporting 
oxygenated blood 


Renal artery BH Vessels transporting 


Renal vein 


deoxygenated blood 


Midneye Bi Vessels involved in 


gas excange 


Lower body 


The pulmonary circuit moves blood from the right side of 
the heart to the lungs and back to the heart. The systemic 
circuit moves blood from the left side of the heart to the 
head and body and returns it to the right side of the heart to 
repeat the cycle. The arrows indicate the direction of blood 
flow, and the colors show the relative levels of oxygen 
concentration. 


Shared Structures 


Different types of blood vessels vary slightly in their structures, but they 
share the same general features. Arteries and arterioles have thicker walls 
than veins and venules because they are closer to the heart and receive 
blood that is surging at a far greater pressure ({link]). Each type of vessel 
has a lumen—a hollow passageway through which blood flows. Arteries 
have smaller lumens than veins, a characteristic that helps to maintain the 
pressure of blood moving through the system. Together, their thicker walls 


and smaller diameters give arterial lumens a more rounded appearance in 
cross section than the lumens of veins. 


Structure of Blood Vessels 
Artery Vein 


= + Tunica externa 


LD i Tunica externa 
Ae 


Tunica media 


S _ Tunica intima 


Smooth muscle 


Internal elastic 
membrane 
Vasa vasorum 
External elastic 
membrane 
Nervi vasorum 
Endothelium 
Elastic fiber 


Endothelium 


(a) Arteries and (b) veins share the same 
general features, but the walls of arteries are 
much thicker because of the higher pressure of 
the blood that flows through them. (c) A 
micrograph shows the relative differences in 
thickness. LM x 160. (Micrograph provided by 
the Regents of the University of Michigan 
Medical School © 2012) 


By the time blood has passed through capillaries and entered venules, the 
pressure initially exerted upon it by heart contractions has diminished. In 
other words, in comparison to arteries, venules and veins withstand a much 
lower pressure from the blood that flows through them. Their walls are 
considerably thinner and their lumens are correspondingly larger in 
diameter, allowing more blood to flow with less vessel resistance. In 
addition, many veins of the body, particularly those of the limbs, contain 
valves that assist the unidirectional flow of blood toward the heart. This is 
critical because blood flow becomes sluggish in the extremities, as a result 
of the lower pressure and the effects of gravity. 


The walls of arteries and veins are largely composed of living cells and 
their products (including collagenous and elastic fibers); the cells require 
nourishment and produce waste. Since blood passes through the larger 
vessels relatively quickly, there is limited opportunity for blood in the 
lumen of the vessel to provide nourishment to or remove waste from the 
vessel’s cells. Further, the walls of the larger vessels are too thick for 
nutrients to diffuse through to all of the cells. Larger arteries and veins 
contain small blood vessels within their walls known as the vasa vasorum 
—literally “vessels of the vessel”—to provide them with this critical 
exchange. Since the pressure within arteries is relatively high, the vasa 
vasorum must function in the outer layers of the vessel (see [link]) or the 
pressure exerted by the blood passing through the vessel would collapse it, 
preventing any exchange from occurring. The lower pressure within veins 
allows the vasa vasorum to be located closer to the lumen. The restriction of 
the vasa vasorum to the outer layers of arteries is thought to be one reason 
that arterial diseases are more common than venous diseases, since its 
location makes it more difficult to nourish the cells of the arteries and 
remove waste products. There are also minute nerves within the walls of 
both types of vessels that control the contraction and dilation of smooth 
muscle. These minute nerves are known as the nervi vasorum. 


Both arteries and veins have the same three distinct tissue layers, called 
tunics (from the Latin term tunica), for the garments first worn by ancient 
Romans; the term tunic is also used for some modem garments. From the 


most interior layer to the outer, these tunics are the tunica intima, the tunica 
media, and the tunica externa (see [link]). [link] compares and contrasts the 


tunics of the arteries and veins. 


Comparison of Tunics in Arteries and Veins 


Arteries 
General Thick walls with small lumens 
appearance Generally appear rounded 


Endothelium usually appears 
wavy due to constriction of 
smooth muscle 

Internal elastic membrane 
present in larger vessels 


Tunica 
intima 


Veins 


Thin walls 
with large 
lumens 
Generally 
appear 
flattened 


Endothelium 
appears 
smooth 
Internal elastic 
membrane 
absent 


Comparison of Tunics in Arteries and Veins 


Tunica 
media 


Tunica 
externa 


Arteries 


Normally the thickest layer in 
arteries 

Smooth muscle cells and elastic 
fibers predominate (the 
proportions of these vary with 
distance from the heart) 
External elastic membrane 
present in larger vessels 


Normally thinner than the tunica 
media in all but the largest 
arteries 

Collagenous and elastic fibers 
Nervi vasorum and vasa 
vasorum present 


Veins 


Normally 
thinner than 
the tunica 
externa 
Smooth 
muscle cells 
and 
collagenous 
fibers 
predominate 
Nervi vasorum 
and vasa 
vasorum 
present 
External 
elastic 
membrane 
absent 


Normally the 
thickest layer 
in veins 
Collagenous 
and smooth 
fibers 
predominate 
Some smooth 
muscle fibers 
Nervi vasorum 
and vasa 
vasorum 
present 


Tunica Intima 


The tunica intima (also called the tunica interna) is composed of epithelial 
and connective tissue layers. Lining the tunica intima is the specialized 
simple squamous epithelium called the endothelium, which is continuous 
throughout the entire vascular system, including the lining of the chambers 
of the heart. Damage to this endothelial lining and exposure of blood to the 
collagenous fibers beneath is one of the primary causes of clot formation. 
Until recently, the endothelium was viewed simply as the boundary between 
the blood in the lumen and the walls of the vessels. Recent studies, 
however, have shown that it is physiologically critical to such activities as 
helping to regulate capillary exchange and altering blood flow. The 
endothelium releases local chemicals called endothelins that can constrict 
the smooth muscle within the walls of the vessel to increase blood pressure. 
Uncompensated overproduction of endothelins may contribute to 
hypertension (high blood pressure) and cardiovascular disease. 


Next to the endothelium is the basement membrane, or basal lamina, that 
effectively binds the endothelium to the connective tissue. The basement 
membrane provides strength while maintaining flexibility, and it is 
permeable, allowing materials to pass through it. The thin outer layer of the 
tunica intima contains a small amount of areolar connective tissue that 
consists primarily of elastic fibers to provide the vessel with additional 
flexibility; it also contains some collagenous fibers to provide additional 
strength. 


In larger arteries, there is also a thick, distinct layer of elastic fibers known 
as the internal elastic membrane (also called the internal elastic lamina) at 
the boundary with the tunica media. Like the other components of the 
tunica intima, the internal elastic membrane provides structure while 
allowing the vessel to stretch. It is permeated with small openings that 
allow exchange of materials between the tunics. The internal elastic 
membrane is not apparent in veins. In addition, many veins, particularly in 
the lower limbs, contain valves formed by sections of thickened 
endothelium that are reinforced with connective tissue, extending into the 
lumen. 


Under the microscope, the lumen and the entire tunica intima of a vein will 
appear smooth, whereas those of an artery will normally appear wavy 
because of the partial constriction of the smooth muscle in the tunica media, 
the next layer of blood vessel walls. 


Tunica Media 


The tunica media is the substantial middle layer of the vessel wall (see 
[link]). It is generally the thickest layer in arteries, and it is much thicker in 
arteries than it is in veins. The tunica media consists of layers of smooth 
muscle supported by connective tissue that is primarily made up of elastic 
fibers, most of which are arranged in circular sheets. Toward the outer 
portion of the tunic, there are also layers of longitudinal muscle. 
Contraction and relaxation of the circular muscles decrease and increase the 
diameter of the vessel lumen, respectively. Specifically in arteries, 
vasoconstriction decreases blood flow as the smooth muscle in the walls of 
the tunica media contracts, making the lumen narrower and increasing 
blood pressure. Similarly, vasodilation increases blood flow as the smooth 
muscle relaxes, allowing the lumen to widen and blood pressure to drop. 
Both vasoconstriction and vasodilation are regulated in part by small 
vascular nerves, known as nervi vasorum, or “nerves of the vessel,” that 
run within the walls of blood vessels. These are generally all sympathetic 
fibers, although some trigger vasodilation and others induce 
vasoconstriction, depending upon the nature of the neurotransmitter and 
receptors located on the target cell. Parasympathetic stimulation does 
trigger vasodilation as well as erection during sexual arousal in the external 
genitalia of both sexes. Nervous control over vessels tends to be more 
generalized than the specific targeting of individual blood vessels. Local 
controls, discussed later, account for this phenomenon. (Seek additional 
content for more information on these dynamic aspects of the autonomic 
nervous system.) Hormones and local chemicals also control blood vessels. 
Together, these neural and chemical mechanisms reduce or increase blood 
flow in response to changing body conditions, from exercise to hydration. 
Regulation of both blood flow and blood pressure is discussed in detail later 
in this chapter. 


The smooth muscle layers of the tunica media are supported by a 
framework of collagenous fibers that also binds the tunica media to the 
inner and outer tunics. Along with the collagenous fibers are large numbers 
of elastic fibers that appear as wavy lines in prepared slides. Separating the 
tunica media from the outer tunica externa in larger arteries is the external 
elastic membrane (also called the external elastic lamina), which also 
appears wavy in Slides. This structure is not usually seen in smaller arteries, 
nor is it seen in veins. 


Tunica Externa 


The outer tunic, the tunica externa (also called the tunica adventitia), is a 
substantial sheath of connective tissue composed primarily of collagenous 
fibers. Some bands of elastic fibers are found here as well. The tunica 
externa in veins also contains groups of smooth muscle fibers. This is 
normally the thickest tunic in veins and may be thicker than the tunica 
media in some larger arteries. The outer layers of the tunica externa are not 
distinct but rather blend with the surrounding connective tissue outside the 
vessel, helping to hold the vessel in relative position. If you are able to 
palpate some of the superficial veins on your upper limbs and try to move 
them, you will find that the tunica externa prevents this. If the tunica 
externa did not hold the vessel in place, any movement would likely result 
in disruption of blood flow. 


Arteries 


An artery is a blood vessel that conducts blood away from the heart. All 
arteries have relatively thick walls that can withstand the high pressure of 
blood ejected from the heart. However, those close to the heart have the 
thickest walls, containing a high percentage of elastic fibers in all three of 
their tunics. This type of artery is known as an elastic artery ((link)). 
Vessels larger than 10 mm in diameter are typically elastic. Their abundant 
elastic fibers allow them to expand, as blood pumped from the ventricles 
passes through them, and then to recoil after the surge has passed. If artery 
walls were rigid and unable to expand and recoil, their resistance to blood 


flow would greatly increase and blood pressure would rise to even higher 
levels, which would in turn require the heart to pump harder to increase the 
volume of blood expelled by each pump (the stroke volume) and maintain 
adequate pressure and flow. Artery walls would have to become even 
thicker in response to this increased pressure. The elastic recoil of the 
vascular wall helps to maintain the pressure gradient that drives the blood 
through the arterial system. An elastic artery is also known as a conducting 
artery, because the large diameter of the lumen enables it to accept a large 
volume of blood from the heart and conduct it to smaller branches. 

Types of Arteries and Arterioles 


Elastic Tunica Muscular Tunica Arteriole Tunica 
artery a— externa artery = = externa = externa 


unica Tunica 
media 


DP Tu nica 
“as 


intima 


Tunica 
media 


b Tunica 


intima 


media 


5 ie Tunica 


intima 


Comparison of the walls of an elastic artery, a muscular artery, 

and an arteriole is shown. In terms of scale, the diameter of an 

arteriole is measured in micrometers compared to millimeters 
for elastic and muscular arteries. 


Farther from the heart, where the surge of blood has dampened, the 
percentage of elastic fibers in an artery’s tunica intima decreases and the 
amount of smooth muscle in its tunica media increases. The artery at this 
point is described as a muscular artery. The diameter of muscular arteries 
typically ranges from 0.1 mm to 10 mm. Their thick tunica media allows 
muscular arteries to play a leading role in vasoconstriction. In contrast, their 
decreased quantity of elastic fibers limits their ability to expand. 
Fortunately, because the blood pressure has eased by the time it reaches 
these more distant vessels, elasticity has become less important. 


Notice that although the distinctions between elastic and muscular arteries 
are important, there is no “line of demarcation” where an elastic artery 
suddenly becomes muscular. Rather, there is a gradual transition as the 


vascular tree repeatedly branches. In turn, muscular arteries branch to 
distribute blood to the vast network of arterioles. For this reason, a 
muscular artery is also known as a distributing artery. 


Arterioles 


An arteriole is a very small artery that leads to a capillary. Arterioles have 
the same three tunics as the larger vessels, but the thickness of each is 
greatly diminished. The critical endothelial lining of the tunica intima is 
intact. The tunica media is restricted to one or two smooth muscle cell 
layers in thickness. The tunica externa remains but is very thin (see [link]). 


With a lumen averaging 30 micrometers or less in diameter, arterioles are 
critical in slowing down—or resisting—blood flow and, thus, causing a 
substantial drop in blood pressure. Because of this, you may see them 
referred to as resistance vessels. The muscle fibers in arterioles are 
normally slightly contracted, causing arterioles to maintain a consistent 
muscle tone—in this case referred to as vascular tone—in a similar manner 
to the muscular tone of skeletal muscle. In reality, all blood vessels exhibit 
vascular tone due to the partial contraction of smooth muscle. The 
importance of the arterioles is that they will be the primary site of both 
resistance and regulation of blood pressure. The precise diameter of the 
lumen of an arteriole at any given moment is determined by neural and 
chemical controls, and vasoconstriction and vasodilation in the arterioles 
are the primary mechanisms for distribution of blood flow. 


Capillaries 


A capillary is a microscopic channel that supplies blood to the tissues 
themselves, a process called perfusion. Exchange of gases and other 
substances occurs in the capillaries between the blood and the surrounding 
cells and their tissue fluid (interstitial fluid). The diameter of a capillary 
lumen ranges from 5—10 micrometers; the smallest are just barely wide 
enough for an erythrocyte to squeeze through. Flow through capillaries is 
often described as microcirculation. 


The wall of a capillary consists of the endothelial layer surrounded by a 
basement membrane with occasional smooth muscle fibers. There is some 
variation in wall structure: In a large capillary, several endothelial cells 
bordering each other may line the lumen; in a small capillary, there may be 
only a single cell layer that wraps around to contact itself. 


For capillaries to function, their walls must be leaky, allowing substances to 
pass through. There are three major types of capillaries, which differ 
according to their degree of “leakiness:” continuous, fenestrated, and 
sinusoid capillaries ({Link]). 


Continuous Capillaries 


The most common type of capillary, the continuous capillary, is found in 
almost all vascularized tissues. Continuous capillaries are characterized by 
a complete endothelial lining with tight junctions between endothelial cells. 
Although a tight junction is usually impermeable and only allows for the 
passage of water and ions, they are often incomplete in capillaries, leaving 
intercellular clefts that allow for exchange of water and other very small 
molecules between the blood plasma and the interstitial fluid. Substances 
that can pass between cells include metabolic products, such as glucose, 
water, and small hydrophobic molecules like gases and hormones, as well 
as various leukocytes. Continuous capillaries not associated with the brain 
are rich in transport vesicles, contributing to either endocytosis or 
exocytosis. Those in the brain are part of the blood-brain barrier. Here, there 
are tight junctions and no intercellular clefts, plus a thick basement 
membrane and astrocyte extensions called end feet; these structures 
combine to prevent the movement of nearly all substances. 

Types of Capillaries 


Continuous Fenestrated Sinusoid 


Endothelial layer 
(tunica intima) 


Incomplete 
basement 
membrane 


Intercellular cleft Fenestrations Intercellular gap 
The three major types of capillaries: continuous, 
fenestrated, and sinusoid. 


Fenestrated Capillaries 


A fenestrated capillary is one that has pores (or fenestrations) in addition 
to tight junctions in the endothelial lining. These make the capillary 
permeable to larger molecules. The number of fenestrations and their 
degree of permeability vary, however, according to their location. 
Fenestrated capillaries are common in the small intestine, which is the 
primary site of nutrient absorption, as well as in the kidneys, which filter 
the blood. They are also found in the choroid plexus of the brain and many 
endocrine structures, including the hypothalamus, pituitary, pineal, and 
thyroid glands. 


Sinusoid Capillaries 


A sinusoid capillary (or sinusoid) is the least common type of capillary. 
Sinusoid capillaries are flattened, and they have extensive intercellular gaps 
and incomplete basement membranes, in addition to intercellular clefts and 
fenestrations. This gives them an appearance not unlike Swiss cheese. 
These very large openings allow for the passage of the largest molecules, 
including plasma proteins and even cells. Blood flow through sinusoids is 


very slow, allowing more time for exchange of gases, nutrients, and wastes. 
Sinusoids are found in the liver and spleen, bone marrow, lymph nodes 
(where they carry lymph, not blood), and many endocrine glands including 
the pituitary and adrenal glands. Without these specialized capillaries, these 
organs would not be able to provide their myriad of functions. For example, 
when bone marrow forms new blood cells, the cells must enter the blood 
supply and can only do so through the large openings of a sinusoid 
capillary; they cannot pass through the small openings of continuous or 
fenestrated capillaries. The liver also requires extensive specialized sinusoid 
capillaries in order to process the materials brought to it by the hepatic 
portal vein from both the digestive tract and spleen, and to release plasma 
proteins into circulation. 


Metarterioles and Capillary Beds 


A metarteriole is a type of vessel that has structural characteristics of both 
an arteriole and a capillary. Slightly larger than the typical capillary, the 
smooth muscle of the tunica media of the metarteriole is not continuous but 
forms rings of smooth muscle (sphincters) prior to the entrance to the 
capillaries. Each metarteriole arises from a terminal arteriole and branches 
to supply blood to a capillary bed that may consist of 10—100 capillaries. 


The precapillary sphincters, circular smooth muscle cells that surround 
the capillary at its origin with the metarteriole, tightly regulate the flow of 
blood from a metarteriole to the capillaries it supplies. Their function is 
critical: If all of the capillary beds in the body were to open simultaneously, 
they would collectively hold every drop of blood in the body and there 
would be none in the arteries, arterioles, venules, veins, or the heart itself. 
Normally, the precapillary sphincters are closed. When the surrounding 
tissues need oxygen and have excess waste products, the precapillary 
sphincters open, allowing blood to flow through and exchange to occur 
before closing once more ((link]). If all of the precapillary sphincters in a 
capillary bed are closed, blood will flow from the metarteriole directly into 
a thoroughfare channel and then into the venous circulation, bypassing the 
capillary bed entirely. This creates what is known as a vascular shunt. In 
addition, an arteriovenous anastomosis may bypass the capillary bed and 
lead directly to the venous system. 


Although you might expect blood flow through a capillary bed to be 
smooth, in reality, it moves with an irregular, pulsating flow. This pattern is 
called vasomotion and is regulated by chemical signals that are triggered in 
response to changes in internal conditions, such as oxygen, carbon dioxide, 
hydrogen ion, and lactic acid levels. For example, during strenuous exercise 
when oxygen levels decrease and carbon dioxide, hydrogen ion, and lactic 
acid levels all increase, the capillary beds in skeletal muscle are open, as 
they would be in the digestive system when nutrients are present in the 
digestive tract. During sleep or rest periods, vessels in both areas are largely 
closed; they open only occasionally to allow oxygen and nutrient supplies 
to travel to the tissues to maintain basic life processes. 


Capillary Bed 
Capillary bed 
eee Capillary 
Arteriole —————————_ Venule 
Precapillary ( ZB | 
sphincter Se SS —— Thoroughfare 


channel 


Metarteriole (serves as 
vascular shunt when 
precapillary sphincters 
are closed) 


Arteriovenous ————____4. i 
anastomosis 


In a capillary bed, arterioles give rise to metarterioles. 
Precapillary sphincters located at the junction of a 
metarteriole with a capillary regulate blood flow. A 
thoroughfare channel connects the metarteriole to a 
venule. An arteriovenous anastomosis, which directly 
connects the arteriole with the venule, is shown at the 
bottom. 


Venules 


A venule is an extremely small vein, generally 8-100 micrometers in 
diameter. Postcapillary venules join multiple capillaries exiting from a 
capillary bed. Multiple venules join to form veins. The walls of venules 
consist of endothelium, a thin middle layer with a few muscle cells and 
elastic fibers, plus an outer layer of connective tissue fibers that constitute a 
very thin tunica externa ({link]). Venules as well as capillaries are the 
primary sites of emigration or diapedesis, in which the white blood cells 
adhere to the endothelial lining of the vessels and then squeeze through 
adjacent cells to enter the tissue fluid. 


Veins 


A vein is a blood vessel that conducts blood toward the heart. Compared to 
arteries, veins are thin-walled vessels with large and irregular lumens (see 
[link]). Because they are low-pressure vessels, larger veins are commonly 
equipped with valves that promote the unidirectional flow of blood toward 
the heart and prevent backflow toward the capillaries caused by the inherent 
low blood pressure in veins as well as the pull of gravity. [link] compares 
the features of arteries and veins. 

Comparison of Veins and Venules 


Large vein 
Tunica externa 


+—— Tunica media 


Garin 
— ® 


Tunica intima 


@ yy Af Smooth muscle cell 
in tunica externa 


Vasa vasorum 


Nervi vasorum 


mate [tunica externa 


+—— Tunica media 


Tunica intima 


Valves 
(closed) 


Tunica externa 
Tunica media 
Tunica intima 


Venule 


Many veins have valves to prevent back 
flow of blood, whereas venules do not. In 
terms of scale, the diameter of a venule is 

measured in micrometers compared to 
millimeters for veins. 


Comparison of Arteries and Veins 


Arteries Veins 
Direction of Conducts Conducts blood toward the 
blood away 
blood flow heart 
from the heart 
General 
Rounded Irregular, often collapsed 
appearance 
Pressure High Low 
ye Il Thick Thin 
thickness 
Higher in 
Relative aaa . : : 
arteries Lower in systemic veins 
oxygen : : : ; 
: Lower in Higher in pulmonary veins 
concentration 
pulmonary 
arteries 
Present most commonly in 
Valves Not present limbs and in veins inferior to 
the heart 
Note: 


Disorders of the... 

Cardiovascular System: Edema and Varicose Veins 

Despite the presence of valves and the contributions of other anatomical 
and physiological adaptations we will cover shortly, over the course of a 
day, some blood will inevitably pool, especially in the lower limbs, due to 
the pull of gravity. Any blood that accumulates in a vein will increase the 


pressure within it, which can then be reflected back into the smaller veins, 
venules, and eventually even the capillaries. Increased pressure will 
promote the flow of fluids out of the capillaries and into the interstitial 
fluid. The presence of excess tissue fluid around the cells leads to a 
condition called edema. 

Most people experience a daily accumulation of tissue fluid, especially if 
they spend much of their work life on their feet (like most health 
professionals). However, clinical edema goes beyond normal swelling and 
requires medical treatment. Edema has many potential causes, including 
hypertension and heart failure, severe protein deficiency, renal failure, and 
many others. In order to treat edema, which is a sign rather than a discrete 
disorder, the underlying cause must be diagnosed and alleviated. 

Varicose Veins 


Varicose veins are commonly 
found in the lower limbs. (credit: 
Thomas Kriese) 


Edema may be accompanied by varicose veins, especially in the superficial 
veins of the legs ([link]). This disorder arises when defective valves allow 
blood to accumulate within the veins, causing them to distend, twist, and 
become visible on the surface of the integument. Varicose veins may occur 
in both sexes, but are more common in women and are often related to 
pregnancy. More than simple cosmetic blemishes, varicose veins are often 
painful and sometimes itchy or throbbing. Without treatment, they tend to 
grow worse over time. The use of support hose, as well as elevating the 
feet and legs whenever possible, may be helpful in alleviating this 
condition. Laser surgery and interventional radiologic procedures can 
reduce the size and severity of varicose veins. Severe cases may require 
conventional surgery to remove the damaged vessels. As there are typically 
redundant circulation patterns, that is, anastomoses, for the smaller and 
more superficial veins, removal does not typically impair the circulation. 
There is evidence that patients with varicose veins suffer a greater risk of 
developing a thrombus or clot. 


Veins as Blood Reservoirs 


In addition to their primary function of returning blood to the heart, veins 
may be considered blood reservoirs, since systemic veins contain 
approximately 64 percent of the blood volume at any given time ([link]). 
Their ability to hold this much blood is due to their high capacitance, that 
is, their capacity to distend (expand) readily to store a high volume of 
blood, even at a low pressure. The large lumens and relatively thin walls of 
veins make them far more distensible than arteries; thus, they are said to be 
Capacitance vessels. 

Distribution of Blood Flow 


Systemic circulation Systemic veins Large veins 
84% 64% 18% 


Large venous networks (liver, bone 
marrow, and integument) 
21% 


Venules and medium-sized veins 
25% 
Systemic arteries Arterioles 
13% 2% 
Muscular arteries 
5% 
Elastic arteries 
4% 
Aorta 
2% 
Systemic capillaries Systemic capillaries 
7% 7% 
Pulmonary circulation Pulmonary veins 
9% 4% 


Pulmonary capillaries 
2% 
se mens a 


3% 


When blood flow needs to be redistributed to other portions of the body, the 
vasomotor center located in the medulla oblongata sends sympathetic 
stimulation to the smooth muscles in the walls of the veins, causing 
constriction—or in this case, venoconstriction. Less dramatic than the 
vasoconstriction seen in smaller arteries and arterioles, venoconstriction 
may be likened to a “stiffening” of the vessel wall. This increases pressure 
on the blood within the veins, speeding its return to the heart. As you will 
note in [link], approximately 21 percent of the venous blood is located in 
venous networks within the liver, bone marrow, and integument. This 
volume of blood is referred to as venous reserve. Through 
venoconstriction, this “reserve” volume of blood can get back to the heart 
more quickly for redistribution to other parts of the circulation. 


Note: 
Career Connection 
Vascular Surgeons and Technicians 


Vascular surgery is a specialty in which the physician deals primarily with 
diseases of the vascular portion of the cardiovascular system. This includes 
repair and replacement of diseased or damaged vessels, removal of plaque 
from vessels, minimally invasive procedures including the insertion of 
venous catheters, and traditional surgery. Following completion of medical 
school, the physician generally completes a 5-year surgical residency 
followed by an additional 1 to 2 years of vascular specialty training. In the 
United States, most vascular surgeons are members of the Society of 
Vascular Surgery. 

Vascular technicians are specialists in imaging technologies that provide 
information on the health of the vascular system. They may also assist 
physicians in treating disorders involving the arteries and veins. This 
profession often overlaps with cardiovascular technology, which would 
also include treatments involving the heart. Although recognized by the 
American Medical Association, there are currently no licensing 
requirements for vascular technicians, and licensing is voluntary. Vascular 
technicians typically have an Associate’s degree or certificate, involving 18 
months to 2 years of training. The United States Bureau of Labor projects 
this profession to grow by 29 percent from 2010 to 2020. 


Note: 

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Visit this site to learn more about vascular surgery. 


Note: 


Visit this site to learn more about vascular technicians. 


Chapter Review 


Blood pumped by the heart flows through a series of vessels known as 
arteries, arterioles, capillaries, venules, and veins before returning to the 
heart. Arteries transport blood away from the heart and branch into smaller 
vessels, forming arterioles. Arterioles distribute blood to capillary beds, the 
sites of exchange with the body tissues. Capillaries lead back to small 
vessels known as venules that flow into the larger veins and eventually back 
to the heart. 


The arterial system is a relatively high-pressure system, so arteries have 
thick walls that appear round in cross section. The venous system is a 
lower-pressure system, containing veins that have larger lumens and thinner 
walls. They often appear flattened. Arteries, arterioles, venules, and veins 
are composed of three tunics known as the tunica intima, tunica media, and 
tunica externa. Capillaries have only a tunica intima layer. The tunica 
intima is a thin layer composed of a simple squamous epithelium known as 
endothelium and a small amount of connective tissue. The tunica media is a 
thicker area composed of variable amounts of smooth muscle and 
connective tissue. It is the thickest layer in all but the largest arteries. The 
tunica externa is primarily a layer of connective tissue, although in veins, it 
also contains some smooth muscle. Blood flow through vessels can be 
dramatically influenced by vasoconstriction and vasodilation in their walls. 


Review Questions 


Exercise: 


Problem:The endothelium is found in the 


a. tunica intima 
b. tunica media 
c. tunica externa 
d. lumen 


Solution: 


A 


Exercise: 


Problem: Nervi vasorum control 


a. vasoconstriction 

b. vasodilation 

c. capillary permeability 

d. both vasoconstriction and vasodilation 


Solution: 


D 
Exercise: 
Problem: 


Closer to the heart, arteries would be expected to have a higher 
percentage of 


a. endothelium 

b. smooth muscle fibers 
c. elastic fibers 

d. collagenous fibers 


Solution: 


@ 


Exercise: 


Problem: Which of the following best describes veins? 


a. thick walled, small lumens, low pressure, lack valves 
b. thin walled, large lumens, low pressure, have valves 

c. thin walled, small lumens, high pressure, have valves 
d. thick walled, large lumens, high pressure, lack valves 


Solution: 


B 
Exercise: 


Problem: 


An especially leaky type of capillary found in the liver and certain 
other tissues is called a 


a. capillary bed 

b. fenestrated capillary 
c. sinusoid capillary 

d. metarteriole 


Solution: 


GC 


Critical Thinking Questions 


Exercise: 


Problem: Arterioles are often referred to as resistance vessels. Why? 


Solution: 


Arterioles receive blood from arteries, which are vessels with a much 
larger lumen. As their own lumen averages just 30 micrometers or less, 
arterioles are critical in slowing down—or resisting—blood flow. The 
arterioles can also constrict or dilate, which varies their resistance, to 
help distribute blood flow to the tissues. 


Exercise: 


Problem: 


Cocaine use causes vasoconstriction. Is this likely to increase or 
decrease blood pressure, and why? 


Solution: 


Vasoconstriction causes the lumens of blood vessels to narrow. This 
increases the pressure of the blood flowing within the vessel. 


Exercise: 


Problem: 


A blood vessel with a few smooth muscle fibers and connective tissue, 
and only a very thin tunica externa conducts blood toward the heart. 
What type of vessel is this? 


Solution: 


This is a venule. 


Glossary 


arteriole 
(also, resistance vessel) very small artery that leads to a capillary 


arteriovenous anastomosis 
short vessel connecting an arteriole directly to a venule and bypassing 
the capillary beds 


artery 
blood vessel that conducts blood away from the heart; may be a 
conducting or distributing vessel 


Capacitance 
ability of a vein to distend and store blood 


Capacitance vessels 
veins 


capillary 
smallest of blood vessels where physical exchange occurs between the 
blood and tissue cells surrounded by interstitial fluid 


capillary bed 
network of 10-100 capillaries connecting arterioles to venules 


continuous capillary 
most common type of capillary, found in virtually all tissues except 
epithelia and cartilage; contains very small gaps in the endothelial 
lining that permit exchange 


elastic artery 
(also, conducting artery) artery with abundant elastic fibers located 
closer to the heart, which maintains the pressure gradient and conducts 
blood to smaller branches 


external elastic membrane 
membrane composed of elastic fibers that separates the tunica media 
from the tunica externa; seen in larger arteries 


fenestrated capillary 
type of capillary with pores or fenestrations in the endothelium that 
allow for rapid passage of certain small materials 


internal elastic membrane 
membrane composed of elastic fibers that separates the tunica intima 
from the tunica media; seen in larger arteries 


lumen 
interior of a tubular structure such as a blood vessel or a portion of the 
alimentary canal through which blood, chyme, or other substances 
travel 


metarteriole 
short vessel arising from a terminal arteriole that branches to supply a 
capillary bed 


microcirculation 
blood flow through the capillaries 


muscular artery 
(also, distributing artery) artery with abundant smooth muscle in the 
tunica media that branches to distribute blood to the arteriole network 


nervi vasorum 
small nerve fibers found in arteries and veins that trigger contraction of 
the smooth muscle in their walls 


perfusion 
distribution of blood into the capillaries so the tissues can be supplied 


precapillary sphincters 
circular rings of smooth muscle that surround the entrance to a 
capillary and regulate blood flow into that capillary 


sinusoid capillary 
rarest type of capillary, which has extremely large intercellular gaps in 
the basement membrane in addition to clefts and fenestrations; found 
in areas such as the bone marrow and liver where passage of large 
molecules occurs 


thoroughfare channel 


continuation of the metarteriole that enables blood to bypass a 
capillary bed and flow directly into a venule, creating a vascular shunt 


tunica externa 
(also, tunica adventitia) outermost layer or tunic of a vessel (except 
capillaries) 


tunica intima 
(also, tunica interna) innermost lining or tunic of a vessel 


tunica media 
middle layer or tunic of a vessel (except capillaries) 


vasa vasorum 
small blood vessels located within the walls or tunics of larger vessels 
that supply nourishment to and remove wastes from the cells of the 
vessels 


vascular shunt 
continuation of the metarteriole and thoroughfare channel that allows 
blood to bypass the capillary beds to flow directly from the arterial to 
the venous circulation 


vasoconstriction 
constriction of the smooth muscle of a blood vessel, resulting in a 
decreased vascular diameter 


vasodilation 
relaxation of the smooth muscle in the wall of a blood vessel, resulting 
in an increased vascular diameter 


vasomotion 
irregular, pulsating flow of blood through capillaries and related 
structures 


vein 
blood vessel that conducts blood toward the heart 


venous reserve 
volume of blood contained within systemic veins in the integument, 
bone marrow, and liver that can be returned to the heart for circulation, 
if needed 


venule 
small vessel leading from the capillaries to veins 


Circulatory Pathways 
By the end of this section, you will be able to: 


e Identify the vessels through which blood travels within the pulmonary 
circuit, beginning from the right ventricle of the heart and ending at the 
left atrium 

e Create a flow chart showing the major systemic arteries through which 
blood travels from the aorta and its major branches, to the most 
significant arteries feeding into the right and left upper and lower 
limbs 

¢ Create a flow chart showing the major systemic veins through which 
blood travels from the feet to the right atrium of the heart 


Virtually every cell, tissue, organ, and system in the body is impacted by the 
circulatory system. This includes the generalized and more specialized 
functions of transport of materials, capillary exchange, maintaining health 
by transporting white blood cells and various immunoglobulins 
(antibodies), hemostasis, regulation of body temperature, and helping to 
maintain acid-base balance. In addition to these shared functions, many 
systems enjoy a unique relationship with the circulatory system. [link] 
summarizes these relationships. 

Interaction of the Circulatory System with Other Body Systems 


Digestive Absorbs nutrients and water; delivers nutrients (except most lipids) to 
liver for processing by hepatic portal vein; provides nutrients essential 
for hematopoiesis and building hemoglobin 


Delivers hormones: atrial natriuretic hormone (peptide) secreted by 
the heart atrial cells to help regulate blood volumes and pressures; 
epinephrine, ANH, angiotensin Il, ADH, and thyroxine to help 
regulate blood pressure; estrogen to promote vascular health in 
women and men 


Carries clotting factors, platelets, and white blood cells for 
hemostasis, fighting infection, and repairing damage; regulates 
temperature by controlling blood flow to the surface, where heat can 
be dissipated; provides some coloration of integument; acts as a 
blood reservoir 


Lymphatic Pes Transports various white blood cells, including those produced by 
lymphatic tissue, and immunoglobulins (antibodies) throughout the 
body to maintain health; carries excess tissue fluid not able to be 
reabsorbed by the vascular capillaries back to the lymphatic system 
for processing 


Provides nutrients and oxygen for contraction; removes lactic acid 
and distributes heat generated by contraction; muscular pumps aid in 
venous return; exercise contributes to cardiovascular health and 
helps to prevent atherosclerosis 


Nervous Produces cerebrospinal fluid (CSF) within choroid plexuses; 
contributes to blood-brain barrier; cardiac and vasomotor centers 
regulate cardiac output and blood flow through vessels via autonomic 
system 


Reproductive : Aids in erection of genitalia in both sexes during sexual arousal; 
transports gonadotropic hormones that regulate reproductive 
functions 


Respiratory Provides blood for critical exchange of gases to carry oxygen needed 
for metabolic reactions and carbon dioxide generated as byproducts 
of these processes 


Skeletal a Provides calcium, phosphate, and other minerals critical for bone 
s matrix; transports hormones regulating buildup and absorption of 
matrix including growth hormone (somatotropin), thyroid hormone, 
calcitonins, and parathyroid hormone; erythropoietin stimulates 
myeloid cell hematopoiesis; some level of protection for select 
vessels by bony structures 


Delivers 20% of resting circulation to kidneys for filtering, 
reabsorption of useful products, and secretion of excesses; regulates 
blood volume and pressure by regulating fluid loss in the form of 
urine and by releasing the enzyme renin that is essential in the 
renin-angiotensin-aldosterone mechanism 


As you learn about the vessels of the systemic and pulmonary circuits, 
notice that many arteries and veins share the same names, parallel one 
another throughout the body, and are very similar on the right and left sides 
of the body. These pairs of vessels will be traced through only one side of 
the body. Where differences occur in branching patterns or when vessels are 
singular, this will be indicated. For example, you will find a pair of femoral 
arteries and a pair of femoral veins, with one vessel on each side of the 
body. In contrast, some vessels closer to the midline of the body, such as the 
aorta, are unique. Moreover, some superficial veins, such as the great 


saphenous vein in the femoral region, have no arterial counterpart. Another 
phenomenon that can make the study of vessels challenging is that names of 
vessels can change with location. Like a street that changes name as it 
passes through an intersection, an artery or vein can change names as it 
passes an anatomical landmark. For example, the left subclavian artery 
becomes the axillary artery as it passes through the body wall and into the 
axillary region, and then becomes the brachial artery as it flows from the 
axillary region into the upper arm (or brachium). You will also find 
examples of anastomoses where two blood vessels that previously branched 
reconnect. Anastomoses are especially common in veins, where they help 
maintain blood flow even when one vessel is blocked or narrowed, although 
there are some important ones in the arteries supplying the brain. 


As you read about circular pathways, notice that there is an occasional, very 
large artery referred to as a trunk, a term indicating that the vessel gives 
rise to several smaller arteries. For example, the celiac trunk gives rise to 
the left gastric, common hepatic, and splenic arteries. 


As you study this section, imagine you are on a “Voyage of Discovery” 
similar to Lewis and Clark’s expedition in 1804—1806, which followed 
rivers and streams through unfamiliar territory, seeking a water route from 
the Atlantic to the Pacific Ocean. You might envision being inside a 
miniature boat, exploring the various branches of the circulatory system. 
This simple approach has proven effective for many students in mastering 
these major circulatory patterns. Another approach that works well for 
many students is to create simple line drawings similar to the ones 
provided, labeling each of the major vessels. It is beyond the scope of this 
text to name every vessel in the body. However, we will attempt to discuss 
the major pathways for blood and acquaint you with the major named 
arteries and veins in the body. Also, please keep in mind that individual 
variations in circulation patterns are not uncommon. 


Note: 


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Visit this site for a brief summary of the arteries. 


Pulmonary Circulation 


Recall that blood returning from the systemic circuit enters the right atrium 
({link]) via the superior and inferior venae cavae and the coronary sinus, 
which drains the blood supply of the heart muscle. These vessels will be 
described more fully later in this section. This blood is relatively low in 
oxygen and relatively high in carbon dioxide, since much of the oxygen has 
been extracted for use by the tissues and the waste gas carbon dioxide was 
picked up to be transported to the lungs for elimination. From the right 
atrium, blood moves into the right ventricle, which pumps it to the lungs for 
gas exchange. This system of vessels is referred to as the pulmonary 
circuit. 


The single vessel exiting the right ventricle is the pulmonary trunk. At the 
base of the pulmonary trunk is the pulmonary semilunar valve, which 
prevents backflow of blood into the right ventricle during ventricular 
diastole. As the pulmonary trunk reaches the superior surface of the heart, it 
curves posteriorly and rapidly bifurcates (divides) into two branches, a left 
and a right pulmonary artery. To prevent confusion between these vessels, 
it is important to refer to the vessel exiting the heart as the pulmonary trunk, 
rather than also calling it a pulmonary artery. The pulmonary arteries in turn 
branch many times within the lung, forming a series of smaller arteries and 
arterioles that eventually lead to the pulmonary capillaries. The pulmonary 
capillaries surround lung structures known as alveoli that are the sites of 
oxygen and carbon dioxide exchange. 


Once gas exchange is completed, oxygenated blood flows from the 
pulmonary capillaries into a series of pulmonary venules that eventually 
lead to a series of larger pulmonary veins. Four pulmonary veins, two on 
the left and two on the right, return blood to the left atrium. At this point, 
the pulmonary circuit is complete. [link] defines the major arteries and 
veins of the pulmonary circuit discussed in the text. 

Pulmonary Circuit 


Ascending aorta Aortic arch 


Superior vena cava Pulmonary trunk 


Right lung Left lung 


Left pulmonary 
arteries 


Left pulmonary 


Right pulmonary wink 


arteries 


Right pulmonary 
veins 


~ Pulmonary 
capillaries 
Inferior vena 


cava Descending 


aorta 


Blood exiting from the right ventricle flows 
into the pulmonary trunk, which bifurcates into 
the two pulmonary arteries. These vessels 
branch to supply blood to the pulmonary 
capillaries, where gas exchange occurs within 
the lung alveoli. Blood returns via the 
pulmonary veins to the left atrium. 


Pulmonary Arteries and Veins 


Vessel Description 


Pulmonary Arteries and Veins 


Vessel Description 
Pulmonary Single large vessel exiting the right ventricle that 
trunk divides to form the right and left pulmonary arteries 
Peionay Left and right vessels that form from the pulmonary 
, a trunk and lead to smaller arterioles and eventually to 

arteries eae fi 

the pulmonary capillaries 

Two sets of paired vessels—one pair on each side— 
Pulmonary that are formed from the small venules, leading away 
veins from the pulmonary capillaries to flow into the left 


atrium 


Overview of Systemic Arteries 


Blood relatively high in oxygen concentration is returned from the 
pulmonary circuit to the left atrium via the four pulmonary veins. From the 
left atrium, blood moves into the left ventricle, which pumps blood into the 
aorta. The aorta and its branches—the systemic arteries—send blood to 
virtually every organ of the body ([link]). 

Systemic Arteries 


Vertebral 
es Right common carotid 


Left common carotid 
Left subclavian 
Axillary 


il Pulmonary trunk 


Right subclavian 


Brachiocephalic trunk 
Aortic arch 


Ascending aorta Br VT . Descending aorta 
SN \) Diaphragm 

Celiac trunk = <h A\ Renal 

= ena 

H Se vant \ 
Brachial P = Superior mesenteric 
/ a 
Radial / | j j We aN Inferior mesenteric 
ii Common iliac 
Ulnar Lh: 
i} y us ae Internal iliac 
External iliac a o 
AY A) 
A 
Palmar AN fea w iS y 
arches ead ait y= Deep 
EAN femoral 
MAW \ ¥ WA 
] F Femoral 
Descending 
\) genicular 


Popliteal 


Posterior tibial 


Anterior tibial 


Fibular 


i 
au 
IN 


Dorsalis pedis 


Plantar arch ae 


The major systemic arteries shown here 
deliver oxygenated blood throughout the 
body. 


The Aorta 


The aorta is the largest artery in the body ([link]). It arises from the left 
ventricle and eventually descends to the abdominal region, where it 
bifurcates at the level of the fourth lumbar vertebra into the two common 
iliac arteries. The aorta consists of the ascending aorta, the aortic arch, and 
the descending aorta, which passes through the diaphragm and a landmark 
that divides into the superior thoracic and inferior abdominal components. 
Arteries originating from the aorta ultimately distribute blood to virtually 
all tissues of the body. At the base of the aorta is the aortic semilunar valve 
that prevents backflow of blood into the left ventricle while the heart is 
relaxing. After exiting the heart, the ascending aorta moves in a superior 
direction for approximately 5 cm and ends at the sternal angle. Following 
this ascent, it reverses direction, forming a graceful arc to the left, called the 
aortic arch. The aortic arch descends toward the inferior portions of the 
body and ends at the level of the intervertebral disk between the fourth and 
fifth thoracic vertebrae. Beyond this point, the descending aorta continues 
close to the bodies of the vertebrae and passes through an opening in the 
diaphragm known as the aortic hiatus. Superior to the diaphragm, the aorta 
is called the thoracic aorta, and inferior to the diaphragm, it is called the 
abdominal aorta. The abdominal aorta terminates when it bifurcates into 
the two common iliac arteries at the level of the fourth lumbar vertebra. See 
[link] for an illustration of the ascending aorta, the aortic arch, and the 
initial segment of the descending aorta plus major branches; [link] 
summarizes the structures of the aorta. 

Aorta 


Right common Left common 
carotid artery carotid artery 


Left subclavian 
artery 


Right subclavian 
artery 


Brachiocephalic 


artery Aortic arch 


Ascending aorta Descending aorta 


Left coronary 


Right coronary artery 


artery 


Thoracic aorta 


Abdominal aorta 


| 


The aorta has distinct regions, including 
the ascending aorta, aortic arch, and the 
descending aorta, which includes the 
thoracic and abdominal regions. 


Components of the Aorta 


Vessel Description 


Components of the Aorta 


Vessel Description 
Largest artery in the body, originating from the left 
ventricle and descending to the abdominal region, 
where it bifurcates into the common iliac arteries at 
Aorta . 
the level of the fourth lumbar vertebra; arteries 
originating from the aorta distribute blood to 
virtually all tissues of the body 
Ascending Initial portion of the aorta, rising superiorly from the 
aorta left ventricle for a distance of approximately 5 cm 
Graceful arc to the left that connects the ascending 
; aorta to the descending aorta; ends at the 
Aortic arch ; ; : 
intervertebral disk between the fourth and fifth 
thoracic vertebrae 
Portion of the aorta that continues inferiorly past the 
Descending ; ws ; ; 
end of the aortic arch; subdivided into the thoracic 
aorta ; 
aorta and the abdominal aorta 
Thoracic Portion of the descending aorta superior to the aortic 
aorta hiatus 
Abdominal Portion of the aorta inferior to the aortic hiatus and 
aorta superior to the common iliac arteries 


Coronary Circulation 


The first vessels that branch from the ascending aorta are the paired 
coronary arteries (see [link]), which arise from two of the three sinuses in 
the ascending aorta just superior to the aortic semilunar valve. These 
sinuses contain the aortic baroreceptors and chemoreceptors critical to 


maintain cardiac function. The left coronary artery arises from the left 
posterior aortic sinus. The right coronary artery arises from the anterior 
aortic sinus. Normally, the right posterior aortic sinus does not give rise to a 
vessel. 


The coronary arteries encircle the heart, forming a ring-like structure that 
divides into the next level of branches that supplies blood to the heart 
tissues. (Seek additional content for more detail on cardiac circulation.) 


Aortic Arch Branches 


There are three major branches of the aortic arch: the brachiocephalic 
artery, the left common carotid artery, and the left subclavian (literally 
“under the clavicle”) artery. As you would expect based upon proximity to 
the heart, each of these vessels is classified as an elastic artery. 


The brachiocephalic artery is located only on the right side of the body; 
there is no corresponding artery on the left. The brachiocephalic artery 
branches into the right subclavian artery and the right common carotid 
artery. The left subclavian and left common carotid arteries arise 
independently from the aortic arch but otherwise follow a similar pattern 
and distribution to the corresponding arteries on the right side (see [link]). 


Each subclavian artery supplies blood to the arms, chest, shoulders, back, 
and central nervous system. It then gives rise to three major branches: the 
internal thoracic artery, the vertebral artery, and the thyrocervical artery. 
The internal thoracic artery, or mammary artery, supplies blood to the 
thymus, the pericardium of the heart, and the anterior chest wall. The 
vertebral artery passes through the vertebral foramen in the cervical 
vertebrae and then through the foramen magnum into the cranial cavity to 
supply blood to the brain and spinal cord. The paired vertebral arteries join 
together to form the large basilar artery at the base of the medulla 
oblongata. This is an example of an anastomosis. The subclavian artery also 
gives rise to the thyrocervical artery that provides blood to the thyroid, the 
cervical region of the neck, and the upper back and shoulder. 


The common carotid artery divides into internal and external carotid 
arteries. The right common carotid artery arises from the brachiocephalic 
artery and the left common carotid artery arises directly from the aortic 
arch. The external carotid artery supplies blood to numerous structures 
within the face, lower jaw, neck, esophagus, and larynx. These branches 
include the lingual, facial, occipital, maxillary, and superficial temporal 
arteries. The internal carotid artery initially forms an expansion known as 
the carotid sinus, containing the carotid baroreceptors and chemoreceptors. 
Like their counterparts in the aortic sinuses, the information provided by 
these receptors is critical to maintaining cardiovascular homeostasis (see 
[link]). 


The internal carotid arteries along with the vertebral arteries are the two 
primary suppliers of blood to the human brain. Given the central role and 
vital importance of the brain to life, it is critical that blood supply to this 
organ remains uninterrupted. Recall that blood flow to the brain is 
remarkably constant, with approximately 20 percent of blood flow directed 
to this organ at any given time. When blood flow is interrupted, even for 
just a few seconds, a transient ischemic attack (TIA), or mini-stroke, may 
occur, resulting in loss of consciousness or temporary loss of neurological 
function. In some cases, the damage may be permanent. Loss of blood flow 
for longer periods, typically between 3 and 4 minutes, will likely produce 
irreversible brain damage or a stroke, also called a cerebrovascular 
accident (CVA). The locations of the arteries in the brain not only provide 
blood flow to the brain tissue but also prevent interruption in the flow of 
blood. Both the carotid and vertebral arteries branch once they enter the 
cranial cavity, and some of these branches form a structure known as the 
arterial circle (or circle of Willis), an anastomosis that is remarkably like a 
traffic circle that sends off branches (in this case, arterial branches to the 
brain). As a rule, branches to the anterior portion of the cerebrum are 
normally fed by the internal carotid arteries; the remainder of the brain 
receives blood flow from branches associated with the vertebral arteries. 


The internal carotid artery continues through the carotid canal of the 
temporal bone and enters the base of the brain through the carotid foramen 
where it gives rise to several branches ([link] and [link]). One of these 
branches is the anterior cerebral artery that supplies blood to the frontal 


lobe of the cerebrum. Another branch, the middle cerebral artery, supplies 
blood to the temporal and parietal lobes, which are the most common sites 
of CVAs. The ophthalmic artery, the third major branch, provides blood to 
the eyes. 


The right and left anterior cerebral arteries join together to form an 
anastomosis called the anterior communicating artery. The initial 
segments of the anterior cerebral arteries and the anterior communicating 
artery form the anterior portion of the arterial circle. The posterior portion 
of the arterial circle is formed by a left and a right posterior 
communicating artery that branches from the posterior cerebral artery, 
which arises from the basilar artery. It provides blood to the posterior 
portion of the cerebrum and brain stem. The basilar artery is an 
anastomosis that begins at the junction of the two vertebral arteries and 
sends branches to the cerebellum and brain stem. It flows into the posterior 
cerebral arteries. [link] summarizes the aortic arch branches, including the 
major branches supplying the brain. 

Arteries Supplying the Head and Neck 


Superficial temporal 


Maxillary 


Occipital \\ 
Internal carotid \ Ss 


CO) \ Facial 
Carotid sinus , 
Lingual 


Vertebral 


External carotid 


The common carotid artery gives rise to the 
external and internal carotid arteries. The 
external carotid artery remains superficial and 
gives rise to many arteries of the head. The 
internal carotid artery first forms the carotid 
sinus and then reaches the brain via the carotid 
canal and carotid foramen, emerging into the 
cranium via the foramen lacerum. The 
vertebral artery branches from the subclavian 
artery and passes through the transverse 
foramen in the cervical vertebrae, entering the 
base of the skull at the vertebral foramen. The 
subclavian artery continues toward the arm as 
the axillary artery. 


Arteries Serving the Brain 


Middle cerebral 


Ophthalmic 


Internal carotid 


Basilar 
Vertebral 


Anterior communicating 


Anterior cerebral 


Posterior communicating 


Posterior cerebral 


This inferior view shows the network of arteries 
serving the brain. The structure is referred to as the 
arterial circle or circle of Willis. 


Aortic Arch Branches and Brain Circulation 


Vessel Description 
Single vessel located on the right side of the 
body; the first vessel branching from the aortic 
Brachiocephalic arch; gives rise to the right subclavian artery and 
artery the right common carotid artery; supplies blood 


to the head, neck, upper limb, and wall of the 
thoracic region 


Aortic Arch Branches and Brain Circulation 


Vessel 


Subclavian 
artery 


Internal 
thoracic artery 


Vertebral artery 


Thyrocervical 
artery 


Common 
carotid artery 


External carotid 
artery 


Description 


The right subclavian artery arises from the 
brachiocephalic artery while the left subclavian 
artery arises from the aortic arch; gives rise to 
the internal thoracic, vertebral, and thyrocervical 
arteries; supplies blood to the arms, chest, 
shoulders, back, and central nervous system 


Also called the mammary artery; arises from the 
subclavian artery; supplies blood to the thymus, 
pericardium of the heart, and anterior chest wall 


Arises from the subclavian artery and passes 
through the vertebral foramen through the 
foramen magnum to the brain; joins with the 
internal carotid artery to form the arterial circle; 
supplies blood to the brain and spinal cord 


Arises from the subclavian artery; supplies 
blood to the thyroid, the cervical region, the 
upper back, and shoulder 


The right common carotid artery arises from the 
brachiocephalic artery and the left common 
carotid artery arises from the aortic arch; each 
gives rise to the extemal and internal carotid 
arteries; supplies the respective sides of the head 
and neck 


Arises from the common carotid artery; supplies 
blood to numerous structures within the face, 
lower jaw, neck, esophagus, and larynx 


Aortic Arch Branches and Brain Circulation 


Vessel 


Internal carotid 
artery 


Arterial circle 
or circle of 
Willis 


Anterior 
cerebral artery 


Middle cerebral 
artery 


Ophthalmic 
artery 


Anterior 
communicating 
artery 


Posterior 
communicating 
artery 


Description 


Arises from the common carotid artery and 
begins with the carotid sinus; goes through the 
carotid canal of the temporal bone to the base of 
the brain; combines with the branches of the 
vertebral artery, forming the arterial circle; 
supplies blood to the brain 


An anastomosis located at the base of the brain 
that ensures continual blood supply; formed 
from the branches of the internal carotid and 
vertebral arteries; supplies blood to the brain 


Arises from the internal carotid artery; supplies 
blood to the frontal lobe of the cerebrum 


Another branch of the internal carotid artery; 
supplies blood to the temporal and parietal lobes 
of the cerebrum 


Branch of the internal carotid artery; supplies 
blood to the eyes 


An anastomosis of the right and left internal 
carotid arteries; supplies blood to the brain 


Branches of the posterior cerebral artery that 
form part of the posterior portion of the arterial 
circle; supplies blood to the brain 


Aortic Arch Branches and Brain Circulation 
Vessel Description 


Branch of the basilar artery that forms a portion 
Posterior of the posterior segment of the arterial circle of 
cerebral artery Willis; supplies blood to the posterior portion of 
the cerebrum and brain stem 


Formed from the fusion of the two vertebral 
arteries; sends branches to the cerebellum, brain 
stem, and the posterior cerebral arteries; the 
main blood supply to the brain stem 


Basilar artery 


Thoracic Aorta and Major Branches 


The thoracic aorta begins at the level of vertebra T5 and continues through 
to the diaphragm at the level of T12, initially traveling within the 
mediastinum to the left of the vertebral column. As it passes through the 
thoracic region, the thoracic aorta gives rise to several branches, which are 
collectively referred to as visceral branches and parietal branches ((link]). 
Those branches that supply blood primarily to visceral organs are known as 
the visceral branches and include the bronchial arteries, pericardial 
arteries, esophageal arteries, and the mediastinal arteries, each named after 
the tissues it supplies. Each bronchial artery (typically two on the left and 
one on the right) supplies systemic blood to the lungs and visceral pleura, in 
addition to the blood pumped to the lungs for oxygenation via the 
pulmonary circuit. The bronchial arteries follow the same path as the 
respiratory branches, beginning with the bronchi and ending with the 
bronchioles. There is considerable, but not total, intermingling of the 
systemic and pulmonary blood at anastomoses in the smaller branches of 
the lungs. This may sound incongruous—that is, the mixing of systemic 
arterial blood high in oxygen with the pulmonary arterial blood lower in 
oxygen—but the systemic vessels also deliver nutrients to the lung tissue 
just as they do elsewhere in the body. The mixed blood drains into typical 


pulmonary veins, whereas the bronchial artery branches remain separate 
and drain into bronchial veins described later. Each pericardial artery 
supplies blood to the pericardium, the esophageal artery provides blood to 
the esophagus, and the mediastinal artery provides blood to the 
mediastinum. The remaining thoracic aorta branches are collectively 
referred to as parietal branches or somatic branches, and include the 
intercostal and superior phrenic arteries. Each intercostal artery provides 
blood to the muscles of the thoracic cavity and vertebral column. The 
superior phrenic artery provides blood to the superior surface of the 
diaphragm. [link] lists the arteries of the thoracic region. 

Arteries of the Thoracic and Abdominal Regions 


ici isn : a 
nterna a = | 


Aortic arch 


Visceral branches of 
the thoracic aorta 


Parietal (somatic) J fi Bronchial 

branches of L \ ; <>) — Esophageal 

thoracic aorta L =~ D> L Mediastinal 
= - 4 Ba __ > —__ aw A 

intercostal a , Sa Saewa es Pericardial 


Superior phrenic 


af 
| 
= XN fl Aortic hiatus 


Inferior phrenic > = y 
Diaoh SS Celiac trunk 
iaphragm —__ 7 x a Yh 7 Left gastric 
Adrenal me Splenic 
; = . aS \ ‘: 
Renal i SJ — Common hepatic 


Gonadal : ; 
Superior mesenteric 
Lumbar ————__4#*_———————— SS 
/ A Abdominal aorta 
Median sacral ————_——______*._ =a 
Common iliac [ = Inferior mesenteric 
Internal iliac 


en iliac 


The thoracic aorta gives rise to the arteries of the 
visceral and parietal branches. 


Arteries of the Thoracic Region 


Vessel 


Visceral 
branches 


Bronchial 
artery 


Pericardial 
artery 


Esophageal 
artery 


Mediastinal 
artery 


Parietal 
branches 


Intercostal 
artery 


Superior 
phrenic 
artery 


Description 


A group of arterial branches of the thoracic aorta; 
supplies blood to the viscera (i.e., organs) of the 
thorax 


Systemic branch from the aorta that provides 
oxygenated blood to the lungs; this blood supply is 
in addition to the pulmonary circuit that brings blood 
for oxygenation 


Branch of the thoracic aorta; supplies blood to the 
pericardium 


Branch of the thoracic aorta; supplies blood to the 
esophagus 


Branch of the thoracic aorta; supplies blood to the 
mediastinum 


Also called somatic branches, a group of arterial 
branches of the thoracic aorta; include those that 
supply blood to the thoracic wall, vertebral column, 
and the superior surface of the diaphragm 


Branch of the thoracic aorta; supplies blood to the 
muscles of the thoracic cavity and vertebral column 


Branch of the thoracic aorta; supplies blood to the 
superior surface of the diaphragm 


Abdominal Aorta and Major Branches 


After crossing through the diaphragm at the aortic hiatus, the thoracic aorta 
is called the abdominal aorta (see [link]). This vessel remains to the left of 
the vertebral column and is embedded in adipose tissue behind the 
peritoneal cavity. It formally ends at approximately the level of vertebra L4, 
where it bifurcates to form the common iliac arteries. Before this division, 
the abdominal aorta gives rise to several important branches. A single 
celiac trunk (artery) emerges and divides into the left gastric artery to 
supply blood to the stomach and esophagus, the splenic artery to supply 
blood to the spleen, and the common hepatic artery, which in turn gives 
rise to the hepatic artery proper to supply blood to the liver, the right 
gastric artery to supply blood to the stomach, the cystic artery to supply 
blood to the gall bladder, and several branches, one to supply blood to the 
duodenum and another to supply blood to the pancreas. Two additional 
single vessels arise from the abdominal aorta. These are the superior and 
inferior mesenteric arteries. The superior mesenteric artery arises 
approximately 2.5 cm after the celiac trunk and branches into several major 
vessels that supply blood to the small intestine (duodenum, jejunum, and 
ileum), the pancreas, and a majority of the large intestine. The inferior 
mesenteric artery supplies blood to the distal segment of the large 
intestine, including the rectum. It arises approximately 5 cm superior to the 
common iliac arteries. 


In addition to these single branches, the abdominal aorta gives rise to 
several significant paired arteries along the way. These include the inferior 
phrenic arteries, the adrenal arteries, the renal arteries, the gonadal arteries, 
and the lumbar arteries. Each inferior phrenic artery is a counterpart of a 
superior phrenic artery and supplies blood to the inferior surface of the 
diaphragm. The adrenal artery supplies blood to the adrenal (suprarenal) 
glands and arises near the superior mesenteric artery. Each renal artery 
branches approximately 2.5 cm inferior to the superior mesenteric arteries 
and supplies a kidney. The right renal artery is longer than the left since the 
aorta lies to the left of the vertebral column and the vessel must travel a 
greater distance to reach its target. Renal arteries branch repeatedly to 
supply blood to the kidneys. Each gonadal artery supplies blood to the 
gonads, or reproductive organs, and is also described as either an ovarian 


artery or a testicular artery (internal spermatic), depending upon the sex of 
the individual. An ovarian artery supplies blood to an ovary, uterine 
(Fallopian) tube, and the uterus, and is located within the suspensory 
ligament of the uterus. It is considerably shorter than a testicular artery, 
which ultimately travels outside the body cavity to the testes, forming one 
component of the spermatic cord. The gonadal arteries arise inferior to the 
renal arteries and are generally retroperitoneal. The ovarian artery continues 
to the uterus where it forms an anastomosis with the uterine artery that 
supplies blood to the uterus. Both the uterine arteries and vaginal arteries, 
which distribute blood to the vagina, are branches of the internal iliac artery. 
The four paired lumbar arteries are the counterparts of the intercostal 
arteries and supply blood to the lumbar region, the abdominal wall, and the 
spinal cord. In some instances, a fifth pair of lumbar arteries emerges from 
the median sacral artery. 


The aorta divides at approximately the level of vertebra L4 into a left and a 
right common iliac artery but continues as a small vessel, the median 
sacral artery, into the sacrum. The common iliac arteries provide blood to 
the pelvic region and ultimately to the lower limbs. They split into external 
and internal iliac arteries approximately at the level of the lumbar-sacral 
articulation. Each internal iliac artery sends branches to the urinary 
bladder, the walls of the pelvis, the external genitalia, and the medial 
portion of the femoral region. In females, they also provide blood to the 
uterus and vagina. The much larger external iliac artery supplies blood to 
each of the lower limbs. [link] shows the distribution of the major branches 
of the aorta into the thoracic and abdominal regions. [link] shows the 
distribution of the major branches of the common iliac arteries. [link] 
summarizes the major branches of the abdominal aorta. 

Major Branches of the Aorta 


UNPAIRED E 
Thoracic 


aorta 


Air passages of 
Bronchial respiratory tract, lung 
arteries tissue 


PAIRED 


Vertebrae, spinal 
Pericardial Pericardium Intercostal cord, back muscles, 
arteries arteries body wall, skin 


Vertebrae, spinal 
Esophageal Esophagus I'elilelmdictiiem! Cord, back muscles, 
arteries arteries body wall, skin, 

diaphragm 


Mediastinal Mediastinal structures 
arteries 


Abdominal 
aorta 


Stomach, 
SiMe r-T ce adjacent ities lalg-tltemg) Diaphragm, inferior 
artery portion of arteries portion of esophagus 
esophagus 


Spleen, 
Splenic stomach, Adrenal Adrenal glands 
artery pancreas arteries 


Liver, stomach, 
gallbladder, Renal Kidneys 
duodenum, arteries 

pancreas 


Common 
hepatic 
artery 


Pancreas, small 
Superior intestine, appendix, Gonadal Testes or ovaries 
Hitt Cdlerlaciall first two-thirds of arteries 

large intestine 


Vetebrae, spinal cord, 
abdominal wall, 
lumbar region 


Inferior Last third of large 
mesenteric artery Bintsun-) 


Common iliac 
arteries 


Median 


sacral 
artery 


The flow chart summarizes the distribution of the 
major branches of the aorta into the thoracic and 
abdominal regions. 


Major Branches of the Iliac Arteries 


Abdominal 
aorta 


Left common 
iliac 

(follows pattern 
similar to right 
common iliac) 


Right common Pelvis and right lower 
iliac limb 


Pelvic muscles, skin, 

Right lower (Uaifeln ata) tel aiiiceacciam viscera of pelvis, 
iliac perineum, gluteal 

region, medial thigh 


Superior 


gluteal Hip muscles, hip joint 


Rectum, anus, 
Internal perineal muscles, 
pudendal external genitalia, 
lateral rotators of hip 


Ilium, hip and thigh 
Obturator muscles, hip joint, 
femoral head 


Lateral Skin and muscles of 
sacral sacrum 


The flow chart summarizes the distribution of the 
major branches of the common iliac arteries into the 
pelvis and lower limbs. The left side follows a similar 
pattern to the right. 


Vessels of the Abdominal Aorta 


Vessel Description 


Vessels of the Abdominal Aorta 


Vessel 


Celiac 
trunk 


Left 
gastric 
artery 


Splenic 
artery 


Common 
hepatic 
artery 


Hepatic 
artery 
proper 


Right 
gastric 
artery 


Cystic 
artery 


Superior 
mesenteric 
artery 


Description 


Also called the celiac artery; a major branch of the 
abdominal aorta; gives rise to the left gastric artery, 
the splenic artery, and the common hepatic artery that 
forms the hepatic artery to the liver, the right gastric 
artery to the stomach, and the cystic artery to the gall 
bladder 


Branch of the celiac trunk; supplies blood to the 
stomach 


Branch of the celiac trunk; supplies blood to the 
spleen 


Branch of the celiac trunk that forms the hepatic 
artery, the right gastric artery, and the cystic artery 


Branch of the common hepatic artery; supplies 
systemic blood to the liver 


Branch of the common hepatic artery; supplies blood 
to the stomach 


Branch of the common hepatic artery; supplies blood 
to the gall bladder 


Branch of the abdominal aorta; supplies blood to the 
small intestine (duodenum, jejunum, and ileum), the 
pancreas, and a majority of the large intestine 


Vessels of the Abdominal Aorta 


Vessel 


Inferior 
mesenteric 
artery 


Inferior 
phrenic 
arteries 


Adrenal 
artery 


Renal 


artery 


Gonadal 
artery 


Ovarian 
artery 


Testicular 
artery 


Lumbar 
arteries 


Common 
iliac artery 


Description 


Branch of the abdominal aorta; supplies blood to the 
distal segment of the large intestine and rectum 


Branches of the abdominal aorta; supply blood to the 
inferior surface of the diaphragm 


Branch of the abdominal aorta; supplies blood to the 
adrenal (suprarenal) glands 


Branch of the abdominal aorta; supplies each kidney 


Branch of the abdominal aorta; supplies blood to the 
gonads or reproductive organs; also described as 
Ovarian arteries or testicular arteries, depending upon 
the sex of the individual 


Branch of the abdominal aorta; supplies blood to 
ovary, uterine (Fallopian) tube, and uterus 


Branch of the abdominal aorta; ultimately travels 
outside the body cavity to the testes and forms one 
component of the spermatic cord 


Branches of the abdominal aorta; supply blood to the 
lumbar region, the abdominal wall, and spinal cord 


Branch of the aorta that leads to the internal and 
external iliac arteries 


Vessels of the Abdominal Aorta 


Vessel Description 

Median 

sacral Continuation of the aorta into the sacrum 
artery 


Branch from the common iliac arteries; supplies 
blood to the urinary bladder, walls of the pelvis, 
extemal genitalia, and the medial portion of the 
femoral region; in females, also provides blood to the 
uterus and vagina 


Internal 
iliac artery 


Branch of the common iliac artery that leaves the 
body cavity and becomes a femoral artery; supplies 
blood to the lower limbs 


External 
iliac artery 


Arteries Serving the Upper Limbs 


As the subclavian artery exits the thorax into the axillary region, it is 
renamed the axillary artery. Although it does branch and supply blood to 
the region near the head of the humerus (via the humeral circumflex 
arteries), the majority of the vessel continues into the upper arm, or 
brachium, and becomes the brachial artery ({link]). The brachial artery 
supplies blood to much of the brachial region and divides at the elbow into 
several smaller branches, including the deep brachial arteries, which 
provide blood to the posterior surface of the arm, and the ulnar collateral 
arteries, which supply blood to the region of the elbow. As the brachial 
artery approaches the coronoid fossa, it bifurcates into the radial and ulnar 
arteries, which continue into the forearm, or antebrachium. The radial 
artery and ulnar artery parallel their namesake bones, giving off smaller 
branches until they reach the wrist, or carpal region. At this level, they fuse 
to form the superficial and deep palmar arches that supply blood to the 
hand, as well as the digital arteries that supply blood to the digits. [link] 


shows the distribution of systemic arteries from the heart into the upper 
limb. [link] summarizes the arteries serving the upper limbs. 
Major Arteries Serving the Thorax and Upper Limb 


Right subclavian 


Axillary i —— 


Y ” ea 
Humeral circumflex —~L Ae 
be 
| i —— 
Deep brachial > TF 


Brachial || 


Ulnar collateral Tf] PS 
ll WA 
Radial ny ee 
Anterior crural 
interosseous 
Ulnar 


Deep palmar arch 
Superficial palmar 


[~ arch 
| [; Digital 


The arteries that supply blood 
to the arms and hands are 
extensions of the subclavian 
arteries. 


Major Arteries of the Upper Limb 


Spinal cord, cervical F 

vertebrae, fuses with left ipifelnis ight Het 
vertebral to form basilar EWS sé] 
artery in cranium 


Left 
vertebral 


common common 
carotid carotid 


Muscles, tissues A 

. 7 ween Right - 5 ; Left 
skin of neck, thyroid = Right Brachiocephalic Left A 
gland, shoulders, diesen ASS ETEY anal mUbclavian pial 
upper back 


Right Skin and muscles of 

pectoral Right chest and abdomen, ae Ap Aortic eile al 
and axilla JEYCIETS mammary gland, thoracic arch craareetete 
muscles pericardium 


Left 
axillary 


Structures 
of the arm (isttliis Left 
and elbow MverHie|| brachial 


Thoracic 


Left aorta 


ventricle 
Radial Ulnar 
side of side of 
forearm forearm 


Digital Digital 
arteries arteries 


The flow chart summarizes the distribution of the major 
arteries from the heart into the upper limb. 


Arteries Serving the Upper Limbs 


Description 


Arteries Serving the Upper Limbs 


Vessel 


Axillary 
artery 


Brachial 
artery 


Radial 
artery 


Ulnar 
artery 


Palmar 
arches 
(superficial 
and deep) 


Description 


Continuation of the subclavian artery as it penetrates 
the body wall and enters the axillary region; supplies 
blood to the region near the head of the humerus 
(humeral circumflex arteries); the majority of the 
vessel continues into the brachium and becomes the 
brachial artery 


Continuation of the axillary artery in the brachium; 
supplies blood to much of the brachial region; gives 
off several smaller branches that provide blood to the 
posterior surface of the arm in the region of the 
elbow; bifurcates into the radial and ulnar arteries at 
the coronoid fossa 


Formed at the bifurcation of the brachial artery; 
parallels the radius; gives off smaller branches until 
it reaches the carpal region where it fuses with the 
ulnar artery to form the superficial and deep palmar 
arches; supplies blood to the lower arm and carpal 
region 


Formed at the bifurcation of the brachial artery; 
parallels the ulna; gives off smaller branches until it 
reaches the carpal region where it fuses with the 
radial artery to form the superficial and deep palmar 
arches; supplies blood to the lower arm and carpal 
region 


Formed from anastomosis of the radial and ulnar 
arteries; supply blood to the hand and digital arteries 


Arteries Serving the Upper Limbs 


Vessel Description 
Digital Formed from the superficial and deep palmar arches; 
arteries supply blood to the digits 


Arteries Serving the Lower Limbs 


The external iliac artery exits the body cavity and enters the femoral region 
of the lower leg ({link]). As it passes through the body wall, it is renamed 
the femoral artery. It gives off several smaller branches as well as the 
lateral deep femoral artery that in turn gives rise to a lateral circumflex 
artery. These arteries supply blood to the deep muscles of the thigh as well 
as ventral and lateral regions of the integument. The femoral artery also 
gives rise to the genicular artery, which provides blood to the region of the 
knee. As the femoral artery passes posterior to the knee near the popliteal 
fossa, it is called the popliteal artery. The popliteal artery branches into the 
anterior and posterior tibial arteries. 


The anterior tibial artery is located between the tibia and fibula, and 
supplies blood to the muscles and integument of the anterior tibial region. 
Upon reaching the tarsal region, it becomes the dorsalis pedis artery, 
which branches repeatedly and provides blood to the tarsal and dorsal 
regions of the foot. The posterior tibial artery provides blood to the 
muscles and integument on the posterior surface of the tibial region. The 
fibular or peroneal artery branches from the posterior tibial artery. It 
bifurcates and becomes the medial plantar artery and lateral plantar 
artery, providing blood to the plantar surfaces. There is an anastomosis 
with the dorsalis pedis artery, and the medial and lateral plantar arteries 
form two arches called the dorsal arch (also called the arcuate arch) and 
the plantar arch, which provide blood to the remainder of the foot and 
toes. [link] shows the distribution of the major systemic arteries in the lower 
limb. [link] summarizes the major systemic arteries discussed in the text. 
Major Arteries Serving the Lower Limb 


External iliac ld = Common iliac 
1 


Internal iliac 


Inguinal ligament Lateral sacral 
Internal pudendal 
Deep femoral Suiaise 


Lateral femoral 
circumflex 


Femoral 


Genicular 


Popliteal 


Dorsalis pedis 


Dorsal arch 


Anterior view 


Right external 
iliac 


Deep femoral 


Lateral femoral 
circumflex 


Femoral 


Genicular 


Popliteal 


Peroneal 


Anterior tibial 
Posterior 
tibial 

Fibular 
Lateral plantar 
Medial plantar 
Plantar arch 


Posterior view 


Major arteries serving the lower limb are shown in 
anterior and posterior views. 


Systemic Arteries of the Lower Limb 


External iliac 


Hip joint, 
femoral head, Deep 


Femoral Thigh 
deep thigh femoral 


muscles 
Decending Knee joint, 
genicular skin of leg 
Adductor Median Lateral . 
muscles, femoral femoral Quadriceps Popliteal Leg and 
obturator circumflex circumflex muscles foot 
muscles, hip joint 


Anterior Posterior f 
tibial tibial AlSOEL 


Dorsalis pedis, 
dorsal arch, Peroneal 
plantar arch 


Dorsal ‘ 
metatarsal, Distal foot, 
dorsal digital [imsaed 


The flow chart summarizes the distribution of the systemic 
arteries from the external iliac artery into the lower limb. 


Arteries Serving the Lower Limbs 


Vessel Description 


Arteries Serving the Lower Limbs 


Vessel 


Femoral 
artery 


Deep 
femoral 
artery 


Lateral 


circumflex 


artery 


Genicular 
artery 


Popliteal 
artery 


Anterior 
tibial 
artery 


Dorsalis 
pedis 
artery 


Posterior 
tibial 
artery 


Description 


Continuation of the external iliac artery after it passes 
through the body cavity; divides into several smaller 
branches, the lateral deep femoral artery, and the 
genicular artery; becomes the popliteal artery as it 
passes posterior to the knee 


Branch of the femoral artery; gives rise to the lateral 
circumflex arteries 


Branch of the deep femoral artery; supplies blood to 
the deep muscles of the thigh and the ventral and 
lateral regions of the integument 


Branch of the femoral artery; supplies blood to the 
region of the knee 


Continuation of the femoral artery posterior to the 
knee; branches into the anterior and posterior tibial 
arteries 


Branches from the popliteal artery; supplies blood to 
the anterior tibial region; becomes the dorsalis pedis 
artery 


Forms from the anterior tibial artery; branches 
repeatedly to supply blood to the tarsal and dorsal 
regions of the foot 


Branches from the popliteal artery and gives rise to 
the fibular or peroneal artery; supplies blood to the 
posterior tibial region 


Arteries Serving the Lower Limbs 


Vessel Description 

Medial Arises from the bifurcation of the posterior tibial 

plantar arteries; supplies blood to the medial plantar surfaces 

artery of the foot 

Lateral Arises from the bifurcation of the posterior tibial 

plantar arteries; supplies blood to the lateral plantar surfaces 

artery of the foot 

Dorsal or Formed from the anastomosis of the dorsalis pedis 

arcuate artery and the medial and plantar arteries; branches 

arch supply the distal portions of the foot and digits 
Formed from the anastomosis of the dorsalis pedis 

Plantar : ; 

aah artery and the medial and plantar arteries; branches 


supply the distal portions of the foot and digits 


Overview of Systemic Veins 


Systemic veins return blood to the right atrium. Since the blood has already 
passed through the systemic capillaries, it will be relatively low in oxygen 
concentration. In many cases, there will be veins draining organs and 
regions of the body with the same name as the arteries that supplied these 
regions and the two often parallel one another. This is often described as a 
“complementary” pattern. However, there is a great deal more variability in 
the venous circulation than normally occurs in the arteries. For the sake of 
brevity and clarity, this text will discuss only the most commonly 
encountered patterns. However, keep this variation in mind when you move 
from the classroom to clinical practice. 


In both the neck and limb regions, there are often both superficial and 
deeper levels of veins. The deeper veins generally correspond to the 
complementary arteries. The superficial veins do not normally have direct 


arterial counterparts, but in addition to returning blood, they also make 
contributions to the maintenance of body temperature. When the ambient 
temperature is warm, more blood is diverted to the superficial veins where 
heat can be more easily dissipated to the environment. In colder weather, 
there is more constriction of the superficial veins and blood is diverted 
deeper where the body can retain more of the heat. 


The “Voyage of Discovery” analogy and stick drawings mentioned earlier 
remain valid techniques for the study of systemic veins, but veins present a 
more difficult challenge because there are numerous anastomoses and 
multiple branches. It is like following a river with many tributaries and 
channels, several of which interconnect. Tracing blood flow through 
arteries follows the current in the direction of blood flow, so that we move 
from the heart through the large arteries and into the smaller arteries to the 
capillaries. From the capillaries, we move into the smallest veins and follow 
the direction of blood flow into larger veins and back to the heart. [link] 
outlines the path of the major systemic veins. 


Note: 


Visit this site for a brief online summary of the veins. 


Major Systemic Veins of the Body 


External jugular 


Subclavian <= 

fi ay > 
Axillary ee } 
Cephalic ——————_f/_// | 
Brachial T \ / J 
Basilic I) ST 
Hepatic if | A 


Median cubital 


Radial 


Median antebrachial 


Ulnar 


Palmar venous 
arches 


Digital 


vA y 
AK 


Popliteal 


Small saphenous ——y ||) 


Fibular 


Plantar venous arch 


Dorsal venous arch 


Internal jugular 
Brachiocephalic 
Superior vena cava 


Intercostal 


Inferior vena cava 


Saas . imvy Renal 
, aa. Gonadal 
Tt ..— Lumbar 


Right and left 
common iliac 


External iliac 


Internal iliac 


Posterior tibial 
Anterior tibial 


The major systemic veins of the body are shown 
here in an anterior view. 


The right atrium receives all of the systemic venous return. Most of the 
blood flows into either the superior vena cava or inferior vena cava. If you 
draw an imaginary line at the level of the diaphragm, systemic venous 
circulation from above that line will generally flow into the superior vena 
cava; this includes blood from the head, neck, chest, shoulders, and upper 
limbs. The exception to this is that most venous blood flow from the 
coronary veins flows directly into the coronary sinus and from there directly 
into the right atrium. Beneath the diaphragm, systemic venous flow enters 
the inferior vena cava, that is, blood from the abdominal and pelvic regions 
and the lower limbs. 


The Superior Vena Cava 


The superior vena cava drains most of the body superior to the diaphragm 
({link]). On both the left and right sides, the subclavian vein forms when 
the axillary vein passes through the body wall from the axillary region. It 
fuses with the external and internal jugular veins from the head and neck to 
form the brachiocephalic vein. Each vertebral vein also flows into the 
brachiocephalic vein close to this fusion. These veins arise from the base of 
the brain and the cervical region of the spinal cord, and flow largely through 
the intervertebral foramina in the cervical vertebrae. They are the 
counterparts of the vertebral arteries. Each internal thoracic vein, also 
known as an internal mammary vein, drains the anterior surface of the chest 
wall and flows into the brachiocephalic vein. 


The remainder of the blood supply from the thorax drains into the azygos 
vein. Each intercostal vein drains muscles of the thoracic wall, each 
esophageal vein delivers blood from the inferior portions of the esophagus, 
each bronchial vein drains the systemic circulation from the lungs, and 
several smaller veins drain the mediastinal region. Bronchial veins carry 
approximately 13 percent of the blood that flows into the bronchial arteries; 
the remainder intermingles with the pulmonary circulation and returns to 
the heart via the pulmonary veins. These veins flow into the azygos vein, 
and with the smaller hemiazygos vein (hemi- = “half”) on the left of the 
vertebral column, drain blood from the thoracic region. The hemiazygos 


vein does not drain directly into the superior vena cava but enters the 
brachiocephalic vein via the superior intercostal vein. 


The azygos vein passes through the diaphragm from the thoracic cavity on 
the right side of the vertebral column and begins in the lumbar region of the 
thoracic cavity. It flows into the superior vena cava at approximately the 
level of T2, making a significant contribution to the flow of blood. It 
combines with the two large left and right brachiocephalic veins to form the 
superior vena Cava. 


[link] summarizes the veins of the thoracic region that flow into the 
superior vena Cava. 
Veins of the Thoracic and Abdominal Regions 


Vertebral 


Internal jugular 


Superior . 
vena cava External jugular 
Subclavian 
iad Brachiocephalic 
Axillary 
Esophageal Bsonaie 
Internal 
werdes Hemiazygos 
Azygos 
Intercostal 
Hepatic 


Inferior vena cava 
Renal Phrenic 


Gonadal 
Adrenal 
Lumbar 


Common iliac 


Internal iliac 
External iliac 


Veins of the thoracic and abdominal regions drain 
blood from the area above the diaphragm, returning it 
to the right atrium via the superior vena cava. 


Veins of the Thoracic Region 


Vessel 


Superior vena 
cava 


Subclavian vein 


Brachiocephalic 
veins 


Vertebral vein 


Description 


Large systemic vein; drains blood from most 
areas superior to the diaphragm; empties into the 
right atrium 


Located deep in the thoracic cavity; formed by 
the axillary vein as it enters the thoracic cavity 
from the axillary region; drains the axillary and 
smaller local veins near the scapular region and 
leads to the brachiocephalic vein 


Pair of veins that form from a fusion of the 
external and internal jugular veins and the 
subclavian vein; subclavian, external and 
internal jugulars, vertebral, and internal thoracic 
veins flow into it; drain the upper thoracic 
region and lead to the superior vena cava 


Arises from the base of the brain and the 
cervical region of the spinal cord; passes 
through the intervertebral foramina in the 
cervical vertebrae; drains smaller veins from the 
cranium, spinal cord, and vertebrae, and leads to 
the brachiocephalic vein; counterpart of the 
vertebral artery 


Veins of the Thoracic Region 
Vessel Description 
Also called internal mammary veins; drain the 


anterior surface of the chest wall and lead to the 
brachiocephalic vein 


Internal 
thoracic veins 


Drains the muscles of the thoracic wall and 
Intercostal vein 
leads to the azygos vein 
Esophageal Drains the inferior portions of the esophagus 
vein and leads to the azygos vein 


Drains the systemic circulation from the lungs 

Bronchial vein . 

and leads to the azygos vein 

Originates in the lumbar region and passes 

through the diaphragm into the thoracic cavity 

on the right side of the vertebral column; drains 

AZzygos vein blood from the intercostal veins, esophageal 
veins, bronchial veins, and other veins draining 
the mediastinal region, and leads to the superior 
vena cava 


Smaller vein complementary to the azygos vein; 
drains the esophageal veins from the esophagus 
and the left intercostal veins, and leads to the 
brachiocephalic vein via the superior intercostal 
vein 


Hemiazygos 
vein 


Veins of the Head and Neck 


Blood from the brain and the superficial facial vein flow into each internal 
jugular vein ({link]). Blood from the more superficial portions of the head, 


scalp, and cranial regions, including the temporal vein and maxillary vein, 
flow into each external jugular vein. Although the external and internal 
jugular veins are separate vessels, there are anastomoses between them 
close to the thoracic region. Blood from the external jugular vein empties 
into the subclavian vein. [link] summarizes the major veins of the head and 
neck. 


Major Veins of the Head and Neck 
Vessel Description 


Parallel to the common carotid artery, which is more or 


Internal less its counterpart, and passes through the jugular 
jugular foramen and canal; primarily drains blood from the 
vein brain, receives the superficial facial vein, and empties 


into the subclavian vein 


Temporal Drains blood from the temporal region and flows into 
vein the external jugular vein 

Maxillary Drains blood from the maxillary region and flows into 
vein the external jugular vein 

External Drains blood from the more superficial portions of the 
jugular head, scalp, and cranial regions, and leads to the 

vein subclavian vein 


Venous Drainage of the Brain 


Circulation to the brain is both critical and complex (see [link]). Many 
smaller veins of the brain stem and the superficial veins of the cerebrum 
lead to larger vessels referred to as intracranial sinuses. These include the 
superior and inferior sagittal sinuses, straight sinus, cavernous sinuses, left 
and right sinuses, the petrosal sinuses, and the occipital sinuses. Ultimately, 
sinuses will lead back to either the inferior jugular vein or vertebral vein. 


Most of the veins on the superior surface of the cerebrum flow into the 
largest of the sinuses, the superior sagittal sinus. It is located midsagittally 
between the meningeal and periosteal layers of the dura mater within the 
falx cerebri and, at first glance in images or models, can be mistaken for the 
subarachnoid space. Most reabsorption of cerebrospinal fluid occurs via the 
chorionic villi (arachnoid granulations) into the superior sagittal sinus. 
Blood from most of the smaller vessels originating from the inferior 
cerebral veins flows into the great cerebral vein and into the straight 
sinus. Other cerebral veins and those from the eye socket flow into the 
cavernous sinus, which flows into the petrosal sinus and then into the 
internal jugular vein. The occipital sinus, sagittal sinus, and straight sinuses 
all flow into the left and right transverse sinuses near the lambdoid suture. 
The transverse sinuses in turn flow into the sigmoid sinuses that pass 
through the jugular foramen and into the internal jugular vein. The internal 
jugular vein flows parallel to the common carotid artery and is more or less 
its counterpart. It empties into the brachiocephalic vein. The veins draining 
the cervical vertebrae and the posterior surface of the skull, including some 
blood from the occipital sinus, flow into the vertebral veins. These parallel 
the vertebral arteries and travel through the transverse foramina of the 
cervical vertebrae. The vertebral veins also flow into the brachiocephalic 
veins. [link] summarizes the major veins of the brain. 

Veins of the Head and Neck 


Superior sagittal sinus 
Inferior sagittal sinus 
Straight sinus 
Occipital 

Temporal 
Cavernous sinus 
Right transverse sinus 


Maxillary 
Occipital sinus 
Facial 
Sigmoid sinus 
Petrosal sinus 
External jugular 


Internal jugular 
Vertebral 


Right subclavian 
Axillary 


Superior vena cava 


This left lateral view shows the veins of the head 
and neck, including the intercranial sinuses. 


Major Veins of the Brain 


Vessel Description 


Major Veins of the Brain 


Vessel 


Superior 
Sagittal 
sinus 


Great 
cerebral 
vein 


Straight 
sinus 


Cavernous 
sinus 


Petrosal 
sinus 


Occipital 
sinus 


Transverse 
sinuses 


Description 


Enlarged vein located midsagittally between the 
meningeal and periosteal layers of the dura mater 
within the falx cerebri; receives most of the blood 
drained from the superior surface of the cerebrum and 
leads to the inferior jugular vein and the vertebral 
vein 


Receives most of the smaller vessels from the inferior 
cerebral veins and leads to the straight sinus 


Enlarged vein that drains blood from the brain; 
receives most of the blood from the great cerebral 
vein and leads to the left or right transverse sinus 


Enlarged vein that receives blood from most of the 
other cerebral veins and the eye socket, and leads to 
the petrosal sinus 


Enlarged vein that receives blood from the cavernous 
sinus and leads into the internal jugular veins 


Enlarged vein that drains the occipital region near the 
falx cerebelli and leads to the left and right transverse 
sinuses, and also the vertebral veins 


Pair of enlarged veins near the lambdoid suture that 
drains the occipital, sagittal, and straight sinuses, and 
leads to the sigmoid sinuses 


Major Veins of the Brain 
Vessel Description 
Enlarged vein that receives blood from the transverse 


sinuses and leads through the jugular foramen to the 
internal jugular vein 


Sigmoid 
sinuses 


Veins Draining the Upper Limbs 


The digital veins in the fingers come together in the hand to form the 
palmar venous arches ((link]). From here, the veins come together to form 
the radial vein, the ulnar vein, and the median antebrachial vein. The radial 
vein and the ulnar vein parallel the bones of the forearm and join together 
at the antebrachium to form the brachial vein, a deep vein that flows into 
the axillary vein in the brachium. 


The median antebrachial vein parallels the ulnar vein, is more medial in 
location, and joins the basilic vein in the forearm. As the basilic vein 
reaches the antecubital region, it gives off a branch called the median 
cubital vein that crosses at an angle to join the cephalic vein. The median 
cubital vein is the most common site for drawing venous blood in humans. 
The basilic vein continues through the arm medially and superficially to the 
axillary vein. 


The cephalic vein begins in the antebrachium and drains blood from the 
superficial surface of the arm into the axillary vein. It is extremely 
superficial and easily seen along the surface of the biceps brachii muscle in 
individuals with good muscle tone and in those without excessive 
subcutaneous adipose tissue in the arms. 


The subscapular vein drains blood from the subscapular region and joins 
the cephalic vein to form the axillary vein. As it passes through the body 
wall and enters the thorax, the axillary vein becomes the subclavian vein. 


Many of the larger veins of the thoracic and abdominal region and upper 
limb are further represented in the flow chart in [link]. [link] summarizes 
the veins of the upper limbs. 

Veins of the Upper Limb 


Subclavian 
Axillary 


Cephalic 


Subscapular 


Brachial 
Basilic 


Median cubital 
Cephalic 
Radial 


Median 
antebrachial 


Why) 
“i 
ik 
Cae 


LAA! 
Al 

} ay Palmar venous arches 
YW 


pr) 
j Deep veins Uy rd / 7 
[i Superficial veins Digital 


This anterior view shows the veins that 
drain the upper limb. 


KEY 


Veins Flowing into the Superior Vena Cava 


Collects blood 

from cranium, |§gile]n3 
spinal cord, vertebral 
vertebrae 


Collects blood ; A 
Right Right Collects blood 

atl ie bath external cule from cranium, 

se Se jugular jugular face, neck 


Right Right Left Left 
subclavian brachiocephalic brachiocephalic subclavian 


F Collects ; 
Right upper Right Left 
limb RUE internal internal Lee roe 
axillary brachial 


(see left limb) Ia peaalens thoracic thoracic 
thoracic wall 


Collects Collects 
wacti Suporte blood from (ULES FS) blood from 
Mediaeunal Le arm medial ESS (sdiElla) arm lateral 


vena cava 
surface surface 


Right Median cubital, 
atrium median 
antebrachial 
Left 
radial 


Hemiazygos 
Esophageal 


Left 
intercostal | 


Digital 


The flow chart summarizes the distribution of the veins 
flowing into the superior vena cava. 


Veins of the Upper Limbs 


Veiselof the UppBekdrmbdon 


Vessel 


Digital 
veins 


Palmar 
venous 
arches 


Radial vein 


Ulnar vein 


Brachial 
vein 


Median 
antebrachial 
vein 


Basilic vein 


Description 


Drain the digits and lead to the palmar arches of the 
hand and dorsal venous arch of the foot 


Drain the hand and digits, and lead to the radial 
vein, ulnar veins, and the median antebrachial vein 


Vein that parallels the radius and radial artery; arises 
from the palmar venous arches and leads to the 
brachial vein 


Vein that parallels the ulna and ulnar artery; arises 
from the palmar venous arches and leads to the 
brachial vein 


Deeper vein of the arm that forms from the radial 
and ulnar veins in the lower arm; leads to the 
axillary vein 


Vein that parallels the ulnar vein but is more medial 
in location; intertwines with the palmar venous 
arches; leads to the basilic vein 


Superficial vein of the arm that arises from the 
median antebrachial vein, intersects with the median 
cubital vein, parallels the ulnar vein, and continues 
into the upper arm; along with the brachial vein, it 
leads to the axillary vein 


Veins of the Upper Limbs 


Vessel Description 


Superficial vessel located in the antecubital region 


Median that links the cephalic vein to the basilic vein in the 

cubital vein form of a v; a frequent site from which to draw 
blood 

Cephalic Superficial vessel in the upper arm; leads to the 

vein axillary vein 

Subscapular Drains blood from the subscapular region and leads 

vein to the axillary vein 

Axillary The major vein in the axillary region; drains the 

vein upper limb and becomes the subclavian vein 


The Inferior Vena Cava 


Other than the small amount of blood drained by the azygos and 
hemiazygos veins, most of the blood inferior to the diaphragm drains into 
the inferior vena cava before it is returned to the heart (see [link]). Lying 
just beneath the parietal peritoneum in the abdominal cavity, the inferior 
vena Cava parallels the abdominal aorta, where it can receive blood from 
abdominal veins. The lumbar portions of the abdominal wall and spinal 
cord are drained by a series of lumbar veins, usually four on each side. The 
ascending lumbar veins drain into either the azygos vein on the right or the 
hemiazygos vein on the left, and return to the superior vena cava. The 
remaining lumbar veins drain directly into the inferior vena cava. 


Blood supply from the kidneys flows into each renal vein, normally the 
largest veins entering the inferior vena cava. A number of other, smaller 


veins empty into the left renal vein. Each adrenal vein drains the adrenal or 
suprarenal glands located immediately superior to the kidneys. The right 
adrenal vein enters the inferior vena cava directly, whereas the left adrenal 
vein enters the left renal vein. 


From the male reproductive organs, each testicular vein flows from the 
scrotum, forming a portion of the spermatic cord. Each ovarian vein drains 
an ovary in females. Each of these veins is generically called a gonadal 
vein. The right gonadal vein empties directly into the inferior vena cava, 
and the left gonadal vein empties into the left renal vein. 


Each side of the diaphragm drains into a phrenic vein; the right phrenic 
vein empties directly into the inferior vena cava, whereas the left phrenic 
vein empties into the left renal vein. Blood supply from the liver drains into 
each hepatic vein and directly into the inferior vena cava. Since the inferior 
vena cava lies primarily to the right of the vertebral column and aorta, the 
left renal vein is longer, as are the left phrenic, adrenal, and gonadal veins. 
The longer length of the left renal vein makes the left kidney the primary 
target of surgeons removing this organ for donation. [link] provides a flow 
chart of the veins flowing into the inferior vena cava. [link] summarizes the 
major veins of the abdominal region. 

Venous Flow into Inferior Vena Cava 


Inferior 


vena cava 


Hepatic 
veins 


Phrenic 


wena Diaphragm 


Gonads (testes or Gonadal 
ovaries) veins 


Adrenal 


F Adrenal glands 
veins 


Spinal cord and body 
wall 


Kidneys 


Right common Left common 
iliac iliac 


Pelvic muscles, skin, 
viscera of pelvis, 
perineum, gluteal region 


Left internal Left external Rater 
iliac iliac limb 


Right lower figile aon Cine! Right internal 
limb iliac iliac 


Lateral 
sacral 
veins 


Superior Internal 
gluteal pudendal 
veins veins 


Obturator 
veins 


The flow chart summarizes veins that deliver blood to the 
inferior vena cava. 


Major Veins of the Abdominal Region 


Vessel 


Inferior 
vena 
cava 


Lumbar 
veins 


Renal 
vein 


Adrenal 


vein 


Testicular 
vein 


Ovarian 
vein 


Gonadal 
vein 


Description 


Large systemic vein that drains blood from areas 
largely inferior to the diaphragm; empties into the right 
atrium 


Series of veins that drain the lumbar portion of the 
abdominal wall and spinal cord; the ascending lumbar 
veins drain into the azygos vein on the right or the 
hemiazygos vein on the left; the remaining lumbar 
veins drain directly into the inferior vena cava 


Largest vein entering the inferior vena cava; drains the 
kidneys and flows into the inferior vena cava 


Drains the adrenal or suprarenal; the right adrenal vein 
enters the inferior vena cava directly and the left 
adrenal vein enters the left renal vein 


Drains the testes and forms part of the spermatic cord; 
the right testicular vein empties directly into the 
inferior vena cava and the left testicular vein empties 
into the left renal vein 


Drains the ovary; the right ovarian vein empties 
directly into the inferior vena cava and the left ovarian 
vein empties into the left renal vein 


Generic term for a vein draining a reproductive organ; 
may be either an ovarian vein or a testicular vein, 
depending on the sex of the individual 


Major Veins of the Abdominal Region 


Vessel Description 
Drains the diaphragm; the right phrenic vein flows into 
Phrenic ener ane 
oh the inferior vena cava and the left phrenic vein empties 
into the left renal vein 
Hepatic Drains systemic blood from the liver and flows into 
vein the inferior vena cava 


Veins Draining the Lower Limbs 


The superior surface of the foot drains into the digital veins, and the inferior 
surface drains into the plantar veins, which flow into a complex series of 
anastomoses in the feet and ankles, including the dorsal venous arch and 
the plantar venous arch ((link]). From the dorsal venous arch, blood 
supply drains into the anterior and posterior tibial veins. The anterior tibial 
vein drains the area near the tibialis anterior muscle and combines with the 
posterior tibial vein and the fibular vein to form the popliteal vein. The 
posterior tibial vein drains the posterior surface of the tibia and joins the 
popliteal vein. The fibular vein drains the muscles and integument in 
proximity to the fibula and also joins the popliteal vein. The small 
saphenous vein located on the lateral surface of the leg drains blood from 
the superficial regions of the lower leg and foot, and flows into to the 
popliteal vein. As the popliteal vein passes behind the knee in the popliteal 
region, it becomes the femoral vein. It is palpable in patients without 
excessive adipose tissue. 


Close to the body wall, the great saphenous vein, the deep femoral vein, and 
the femoral circumflex vein drain into the femoral vein. The great 
saphenous vein is a prominent surface vessel located on the medial surface 
of the leg and thigh that collects blood from the superficial portions of these 
areas. The deep femoral vein, as the name suggests, drains blood from the 
deeper portions of the thigh. The femoral circumflex vein forms a loop 


around the femur just inferior to the trochanters and drains blood from the 
areas in proximity to the head and neck of the femur. 


As the femoral vein penetrates the body wall from the femoral portion of 
the upper limb, it becomes the external iliac vein, a large vein that drains 
blood from the leg to the common iliac vein. The pelvic organs and 
integument drain into the internal iliac vein, which forms from several 
smaller veins in the region, including the umbilical veins that run on either 
side of the bladder. The external and internal iliac veins combine near the 
inferior portion of the sacroiliac joint to form the common iliac vein. In 
addition to blood supply from the external and internal iliac veins, the 
middle sacral vein drains the sacral region into the common iliac vein. 
Similar to the common iliac arteries, the common iliac veins come together 
at the level of L5 to form the inferior vena cava. 


[link] is a flow chart of veins flowing into the lower limb. [link] 
summarizes the major veins of the lower limbs. 
Major Veins Serving the Lower Limbs 


Common iliac 


External iliac 
| Internal iliac 
External iliac = 42 Gluteal Internal 
; Lateral sacral pudendal Gluteal 
Internal pudendal Obturator 
Obturator Femoral 
Femoral 
Deep femoral Deep femoral 
Femoral Femoral 
circumflex circumflex 


Femoral 
Femoral 


Great saphenous Great 


saphenous 


Popliteal 
Popliteal 


Small saphenous Small 
Anterior tibial eppnenGys 


Posterior tibial Anterior tibial 


Posterior 
Fibular tibial 


Fibular 
Lateral plantar 


Medial plantar 


Plantar venous 


Dorsal venous arch 
arch 


Digital 


Anterior view Posterior view 


Anterior and posterior views show the major veins that 
drain the lower limb into the inferior vena cava. 


Major Veins of the Lower Limb 


External iliac 


Collects blood from 
Collects blood Deep Eemoral Great the superficial 
from the thigh femoral EY-)9)il-JaleltESa) veins of the lower 


limb 


Collects blood 
from superficial Small 

veins of the leg saphenous 
and foot 


Popliteal 


Anterior Posterior 


tibial tibial AES 


Lateral and medial plantar, 
dorsal arch, plantar arch 


Metatarsal, Distal foot, 
digital toes 


The flow chart summarizes venous flow from the 
lower limb. 


Veins of the Lower Limbs 


Vessel 


Plantar 
veins 


Dorsal 
venous 
arch 


Plantar 
venous 
arch 


Anterior 
tibial vein 


Posterior 
tibial vein 


Fibular 
vein 


Small 
saphenous 
vein 


Popliteal 
vein 


Description 


Drain the foot and flow into the plantar venous arch 


Drains blood from digital veins and vessels on the 
superior surface of the foot 


Formed from the plantar veins; flows into the anterior 
and posterior tibial veins through anastomoses 


Formed from the dorsal venous arch; drains the area 
near the tibialis anterior muscle and flows into the 
popliteal vein 


Formed from the dorsal venous arch; drains the area 
near the posterior surface of the tibia and flows into 
the popliteal vein 


Drains the muscles and integument near the fibula 
and flows into the popliteal vein 


Located on the lateral surface of the leg; drains blood 
from the superficial regions of the lower leg and foot, 
and flows into the popliteal vein 


Drains the region behind the knee and forms from the 
fusion of the fibular, anterior, and posterior tibial 
veins; flows into the femoral vein 


Veins of the Lower Limbs 


Vessel 


Great 
saphenous 
vein 


Deep 
femoral 
vein 


Femoral 
circumflex 
vein 


Femoral 
vein 


External 
iliac vein 


Internal 
iliac vein 


Middle 
sacral vein 


Description 


Prominent surface vessel located on the medial 
surface of the leg and thigh; drains the superficial 
portions of these areas and flows into the femoral 
vein 


Drains blood from the deeper portions of the thigh 
and flows into the femoral vein 


Forms a loop around the femur just inferior to the 
trochanters; drains blood from the areas around the 
head and neck of the femur; flows into the femoral 
vein 


Drains the upper leg; receives blood from the great 
saphenous vein, the deep femoral vein, and the 
femoral circumflex vein; becomes the external iliac 
vein when it crosses the body wall 


Formed when the femoral vein passes into the body 
cavity; drains the legs and flows into the common 
iliac vein 


Drains the pelvic organs and integument; formed 
from several smaller veins in the region; flows into 
the common iliac vein 


Drains the sacral region and flows into the left 
common iliac vein 


Veins of the Lower Limbs 
Vessel Description 


Flows into the inferior vena cava at the level of L5; 
the left common iliac vein drains the sacral region; 
formed from the union of the external and internal 

iliac veins near the inferior portion of the sacroiliac 
joint 


Common 
iliac vein 


Hepatic Portal System 


The liver is a complex biochemical processing plant. It packages nutrients 
absorbed by the digestive system; produces plasma proteins, clotting 
factors, and bile; and disposes of worn-out cell components and waste 
products. Instead of entering the circulation directly, absorbed nutrients and 
certain wastes (for example, materials produced by the spleen) travel to the 
liver for processing. They do so via the hepatic portal system ((Link)). 
Portal systems begin and end in capillaries. In this case, the initial 
capillaries from the stomach, small intestine, large intestine, and spleen lead 
to the hepatic portal vein and end in specialized capillaries within the liver, 
the hepatic sinusoids. You saw the only other portal system with the 
hypothalamic-hypophyseal portal vessel in the endocrine chapter. 


The hepatic portal system consists of the hepatic portal vein and the veins 
that drain into it. The hepatic portal vein itself is relatively short, beginning 
at the level of L2 with the confluence of the superior mesenteric and splenic 
veins. It also receives branches from the inferior mesenteric vein, plus the 
splenic veins and all their tributaries. The superior mesenteric vein receives 
blood from the small intestine, two-thirds of the large intestine, and the 
stomach. The inferior mesenteric vein drains the distal third of the large 
intestine, including the descending colon, the sigmoid colon, and the 
rectum. The splenic vein is formed from branches from the spleen, 
pancreas, and portions of the stomach, and the inferior mesenteric vein. 
After its formation, the hepatic portal vein also receives branches from the 
gastric veins of the stomach and cystic veins from the gall bladder. The 


hepatic portal vein delivers materials from these digestive and circulatory 
organs directly to the liver for processing. 


Because of the hepatic portal system, the liver receives its blood supply 
from two different sources: from normal systemic circulation via the hepatic 
artery and from the hepatic portal vein. The liver processes the blood from 
the portal system to remove certain wastes and excess nutrients, which are 
stored for later use. This processed blood, as well as the systemic blood that 
came from the hepatic artery, exits the liver via the right, left, and middle 
hepatic veins, and flows into the inferior vena cava. Overall systemic blood 
composition remains relatively stable, since the liver is able to metabolize 
the absorbed digestive components. 

Hepatic Portal System 
Hepatic portal 


Cystic ey 
Gall bladder 


Superior mesenteric 
Gastro-omental () 


Right gastric 


Splenic 
Gastroepiploic 


. Pancreatic 
Pancreaticoduodenal 


Middle colic 


Right colic cis ~~ Inferior mesenteric 
i X Left colic 
Ileocolic J > A 


Intestinal ( on s 
xeeyl 


The liver receives blood from the normal systemic 
circulation via the hepatic artery. It also receives and 
processes blood from other organs, delivered via the 
veins of the hepatic portal system. All blood exits the 
liver via the hepatic vein, which delivers the blood to 
the inferior vena cava. (Different colors are used to 
help distinguish among the different vessels in the 
system.) 


Sigmoid 


Superior rectal 


Chapter Review 


The right ventricle pumps oxygen-depleted blood into the pulmonary trunk 
and right and left pulmonary arteries, which carry it to the right and left 
lungs for gas exchange. Oxygen-rich blood is transported by pulmonary 
veins to the left atrium. The left ventricle pumps this blood into the aorta. 
The main regions of the aorta are the ascending aorta, aortic arch, and 
descending aorta, which is further divided into the thoracic and abdominal 
aorta. The coronary arteries branch from the ascending aorta. After 
oxygenating tissues in the capillaries, systemic blood is returned to the right 
atrium from the venous system via the superior vena cava, which drains 
most of the veins superior to the diaphragm, the inferior vena cava, which 
drains most of the veins inferior to the diaphragm, and the coronary veins 
via the coronary sinus. The hepatic portal system carries blood to the liver 
for processing before it enters circulation. Review the figures provided in 
this section for circulation of blood through the blood vessels. 


Review Questions 


Exercise: 


Problem:The coronary arteries branch off of the 


a. aortic valve 

b. ascending aorta 
c. aortic arch 

d. thoracic aorta 


Solution: 


B 


Exercise: 


Problem: Which of the following statements is true? 


a. The left and right common carotid arteries both branch off of the 
brachiocephalic trunk. 

b. The brachial artery is the distal branch of the axillary artery. 

c. The radial and ulnar arteries join to form the palmar arch. 

d. All of the above are true. 


Solution: 


C 
Exercise: 


Problem: 


Arteries serving the stomach, pancreas, and liver all branch from the 


a. Superior mesenteric artery 
b. inferior mesenteric artery 
c. celiac trunk 

d. splenic artery 


Solution: 
C 
Exercise: 
Problem:The right and left brachiocephalic veins 


a. drain blood from the right and left internal jugular veins 
b. drain blood from the right and left subclavian veins 

c. drain into the superior vena cava 

d. all of the above are true 


Solution: 


D 
Exercise: 


Problem: 


The hepatic portal system delivers blood from the digestive organs to 
the 


a. liver 

b. hypothalamus 
c. spleen 

d. left atrium 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: 


Identify the ventricle of the heart that pumps oxygen-depleted blood 
and the arteries of the body that carry oxygen-depleted blood. 


Solution: 


The right ventricle of the heart pumps oxygen-depleted blood to the 
pulmonary arteries. 


Exercise: 


Problem:What organs do the gonadal veins drain? 


Solution: 


The gonadal veins drain the testes in males and the ovaries in females. 
Exercise: 


Problem: 


What arteries play the leading roles in supplying blood to the brain? 


Solution: 


The internal carotid arteries and the vertebral arteries provide most of 
the brain’s blood supply. 


Glossary 


abdominal aorta 
portion of the aorta inferior to the aortic hiatus and superior to the 
common iliac arteries 


adrenal artery 
branch of the abdominal aorta; supplies blood to the adrenal 
(suprarenal) glands 


adrenal vein 
drains the adrenal or suprarenal glands that are immediately superior to 
the kidneys; the right adrenal vein enters the inferior vena cava directly 
and the left adrenal vein enters the left renal vein 


anterior cerebral artery 
arises from the internal carotid artery; supplies the frontal lobe of the 
cerebrum 


anterior communicating artery 
anastomosis of the right and left internal carotid arteries; supplies 
blood to the brain 


anterior tibial artery 


branches from the popliteal artery; supplies blood to the anterior tibial 
region; becomes the dorsalis pedis artery 


anterior tibial vein 
forms from the dorsal venous arch; drains the area near the tibialis 
anterior muscle and leads to the popliteal vein 


aorta 
largest artery in the body, originating from the left ventricle and 
descending to the abdominal region where it bifurcates into the 
common iliac arteries at the level of the fourth lumbar vertebra; 
arteries originating from the aorta distribute blood to virtually all 
tissues of the body 


aortic arch 
arc that connects the ascending aorta to the descending aorta; ends at 
the intervertebral disk between the fourth and fifth thoracic vertebrae 


aortic hiatus 
opening in the diaphragm that allows passage of the thoracic aorta into 
the abdominal region where it becomes the abdominal aorta 


arterial circle 
(also, circle of Willis) anastomosis located at the base of the brain that 
ensures continual blood supply; formed from branches of the internal 
carotid and vertebral arteries; supplies blood to the brain 


ascending aorta 
initial portion of the aorta, rising from the left ventricle for a distance 
of approximately 5 cm 


axillary artery 
continuation of the subclavian artery as it penetrates the body wall and 
enters the axillary region; supplies blood to the region near the head of 
the humerus (humeral circumflex arteries); the majority of the vessel 
continues into the brachium and becomes the brachial artery 


axillary vein 


major vein in the axillary region; drains the upper limb and becomes 
the subclavian vein 


azygos vein 
originates in the lumbar region and passes through the diaphragm into 
the thoracic cavity on the right side of the vertebral column; drains 
blood from the intercostal veins, esophageal veins, bronchial veins, 
and other veins draining the mediastinal region; leads to the superior 
vena cava 


basilar artery 
formed from the fusion of the two vertebral arteries; sends branches to 
the cerebellum, brain stem, and the posterior cerebral arteries; the main 
blood supply to the brain stem 


basilic vein 
superficial vein of the arm that arises from the palmar venous arches, 
intersects with the median cubital vein, parallels the ulnar vein, and 
continues into the upper arm; along with the brachial vein, it leads to 
the axillary vein 


brachial artery 
continuation of the axillary artery in the brachium; supplies blood to 
much of the brachial region; gives off several smaller branches that 
provide blood to the posterior surface of the arm in the region of the 
elbow; bifurcates into the radial and ulnar arteries at the coronoid fossa 


brachial vein 
deeper vein of the arm that forms from the radial and ulnar veins in the 
lower arm; leads to the axillary vein 


brachiocephalic artery 
single vessel located on the right side of the body; the first vessel 
branching from the aortic arch; gives rise to the right subclavian artery 
and the right common carotid artery; supplies blood to the head, neck, 
upper limb, and wall of the thoracic region 


brachiocephalic vein 


one of a pair of veins that form from a fusion of the external and 
internal jugular veins and the subclavian vein; subclavian, external and 
internal jugulars, vertebral, and internal thoracic veins lead to it; drains 
the upper thoracic region and flows into the superior vena cava 


bronchial artery 
systemic branch from the aorta that provides oxygenated blood to the 
lungs in addition to the pulmonary circuit 


bronchial vein 
drains the systemic circulation from the lungs and leads to the azygos 
vein 


cavernous sinus 
enlarged vein that receives blood from most of the other cerebral veins 
and the eye socket, and leads to the petrosal sinus 


celiac trunk 
(also, celiac artery) major branch of the abdominal aorta; gives rise to 
the left gastric artery, the splenic artery, and the common hepatic artery 
that forms the hepatic artery to the liver, the right gastric artery to the 
stomach, and the cystic artery to the gall bladder 


cephalic vein 
superficial vessel in the upper arm; leads to the axillary vein 


cerebrovascular accident (CVA) 
blockage of blood flow to the brain; also called a stroke 


circle of Willis 
(also, arterial circle) anastomosis located at the base of the brain that 
ensures continual blood supply; formed from branches of the internal 
carotid and vertebral arteries; supplies blood to the brain 


common carotid artery 
right common carotid artery arises from the brachiocephalic artery, and 
the left common carotid arises from the aortic arch; gives rise to the 


external and internal carotid arteries; supplies the respective sides of 
the head and neck 


common hepatic artery 
branch of the celiac trunk that forms the hepatic artery, the right gastric 
artery, and the cystic artery 


common iliac artery 
branch of the aorta that leads to the internal and external iliac arteries 


common iliac vein 
one of a pair of veins that flows into the inferior vena cava at the level 
of L5; the left common iliac vein drains the sacral region; divides into 
external and internal iliac veins near the inferior portion of the 
sacroiliac joint 


cystic artery 
branch of the common hepatic artery; supplies blood to the gall 
bladder 


deep femoral artery 
branch of the femoral artery; gives rise to the lateral circumflex 
arteries 


deep femoral vein 
drains blood from the deeper portions of the thigh and leads to the 
femoral vein 


descending aorta 
portion of the aorta that continues downward past the end of the aortic 
arch; subdivided into the thoracic aorta and the abdominal aorta 


digital arteries 
formed from the superficial and deep palmar arches; supply blood to 
the digits 


digital veins 


drain the digits and feed into the palmar arches of the hand and dorsal 
venous arch of the foot 


dorsal arch 
(also, arcuate arch) formed from the anastomosis of the dorsalis pedis 
artery and medial and plantar arteries; branches supply the distal 
portions of the foot and digits 


dorsal venous arch 
drains blood from digital veins and vessels on the superior surface of 
the foot 


dorsalis pedis artery 
forms from the anterior tibial artery; branches repeatedly to supply 
blood to the tarsal and dorsal regions of the foot 


esophageal artery 
branch of the thoracic aorta; supplies blood to the esophagus 


esophageal vein 
drains the inferior portions of the esophagus and leads to the azygos 
vein 


external carotid artery 
arises from the common carotid artery; supplies blood to numerous 
structures within the face, lower jaw, neck, esophagus, and larynx 


external iliac artery 
branch of the common iliac artery that leaves the body cavity and 
becomes a femoral artery; supplies blood to the lower limbs 


external iliac vein 
formed when the femoral vein passes into the body cavity; drains the 
legs and leads to the common iliac vein 


external jugular vein 
one of a pair of major veins located in the superficial neck region that 
drains blood from the more superficial portions of the head, scalp, and 


cranial regions, and leads to the subclavian vein 


femoral artery 
continuation of the external iliac artery after it passes through the body 
cavity; divides into several smaller branches, the lateral deep femoral 
artery, and the genicular artery; becomes the popliteal artery as it 
passes posterior to the knee 


femoral circumflex vein 
forms a loop around the femur just inferior to the trochanters; drains 
blood from the areas around the head and neck of the femur; leads to 
the femoral vein 


femoral vein 
drains the upper leg; receives blood from the great saphenous vein, the 
deep femoral vein, and the femoral circumflex vein; becomes the 
external iliac vein when it crosses the body wall 


fibular vein 
drains the muscles and integument near the fibula and leads to the 
popliteal vein 


genicular artery 
branch of the femoral artery; supplies blood to the region of the knee 


gonadal artery 
branch of the abdominal aorta; supplies blood to the gonads or 
reproductive organs; also described as ovarian arteries or testicular 
arteries, depending upon the sex of the individual 


gonadal vein 
generic term for a vein draining a reproductive organ; may be either an 
ovarian vein or a testicular vein, depending on the sex of the individual 


great cerebral vein 
receives most of the smaller vessels from the inferior cerebral veins 
and leads to the straight sinus 


great saphenous vein 
prominent surface vessel located on the medial surface of the leg and 
thigh; drains the superficial portions of these areas and leads to the 
femoral vein 


hemiazygos vein 
smaller vein complementary to the azygos vein; drains the esophageal 
veins from the esophagus and the left intercostal veins, and leads to the 
brachiocephalic vein via the superior intercostal vein 


hepatic artery proper 
branch of the common hepatic artery; supplies systemic blood to the 
liver 


hepatic portal system 
specialized circulatory pathway that carries blood from digestive 
organs to the liver for processing before being sent to the systemic 
circulation 


hepatic vein 
drains systemic blood from the liver and flows into the inferior vena 
cava 


inferior mesenteric artery 
branch of the abdominal aorta; supplies blood to the distal segment of 
the large intestine and rectum 


inferior phrenic artery 
branch of the abdominal aorta; supplies blood to the inferior surface of 
the diaphragm 


inferior vena cava 
large systemic vein that drains blood from areas largely inferior to the 
diaphragm; empties into the right atrium 


intercostal artery 
branch of the thoracic aorta; supplies blood to the muscles of the 
thoracic cavity and vertebral column 


intercostal vein 
drains the muscles of the thoracic wall and leads to the azygos vein 


internal carotid artery 
arises from the common carotid artery and begins with the carotid 
sinus; goes through the carotid canal of the temporal bone to the base 
of the brain; combines with branches of the vertebral artery forming 
the arterial circle; supplies blood to the brain 


internal iliac artery 
branch from the common iliac arteries; supplies blood to the urinary 
bladder, walls of the pelvis, external genitalia, and the medial portion 
of the femoral region; in females, also provide blood to the uterus and 
vagina 


internal iliac vein 
drains the pelvic organs and integument; formed from several smaller 
veins in the region; leads to the common iliac vein 


internal jugular vein 
one of a pair of major veins located in the neck region that passes 
through the jugular foramen and canal, flows parallel to the common 
carotid artery that is more or less its counterpart; primarily drains 
blood from the brain, receives the superficial facial vein, and empties 
into the subclavian vein 


internal thoracic artery 
(also, mammary artery) arises from the subclavian artery; supplies 
blood to the thymus, pericardium of the heart, and the anterior chest 
wall 


internal thoracic vein 
(also, internal mammary vein) drains the anterior surface of the chest 
wall and leads to the brachiocephalic vein 


lateral circumflex artery 
branch of the deep femoral artery; supplies blood to the deep muscles 
of the thigh and the ventral and lateral regions of the integument 


lateral plantar artery 
arises from the bifurcation of the posterior tibial arteries; supplies 
blood to the lateral plantar surfaces of the foot 


left gastric artery 
branch of the celiac trunk; supplies blood to the stomach 


lumbar arteries 
branches of the abdominal aorta; supply blood to the lumbar region, 
the abdominal wall, and spinal cord 


lumbar veins 
drain the lumbar portion of the abdominal wall and spinal cord; the 
superior lumbar veins drain into the azygos vein on the right or the 
hemiazygos vein on the left; blood from these vessels is returned to the 
superior vena cava rather than the inferior vena cava 


maxillary vein 
drains blood from the maxillary region and leads to the external 
jugular vein 


medial plantar artery 
arises from the bifurcation of the posterior tibial arteries; supplies 
blood to the medial plantar surfaces of the foot 


median antebrachial vein 
vein that parallels the ulnar vein but is more medial in location; 
intertwines with the palmar venous arches 


median cubital vein 
superficial vessel located in the antecubital region that links the 
cephalic vein to the basilic vein in the form of a v; a frequent site for a 
blood draw 


median sacral artery 
continuation of the aorta into the sacrum 


mediastinal artery 


branch of the thoracic aorta; supplies blood to the mediastinum 


middle cerebral artery 
another branch of the internal carotid artery; supplies blood to the 
temporal and parietal lobes of the cerebrum 


middle sacral vein 
drains the sacral region and leads to the left common iliac vein 


occipital sinus 
enlarged vein that drains the occipital region near the falx cerebelli and 
flows into the left and right transverse sinuses, and also into the 
vertebral veins 


ophthalmic artery 
branch of the internal carotid artery; supplies blood to the eyes 


ovarian artery 
branch of the abdominal aorta; supplies blood to the ovary, uterine 
(Fallopian) tube, and uterus 


ovarian vein 
drains the ovary; the right ovarian vein leads to the inferior vena cava 
and the left ovarian vein leads to the left renal vein 


palmar arches 
superficial and deep arches formed from anastomoses of the radial and 
ulnar arteries; supply blood to the hand and digital arteries 


palmar venous arches 
drain the hand and digits, and feed into the radial and ulnar veins 


parietal branches 
(also, somatic branches) group of arterial branches of the thoracic 
aorta; includes those that supply blood to the thoracic cavity, vertebral 
column, and the superior surface of the diaphragm 


pericardial artery 


branch of the thoracic aorta; supplies blood to the pericardium 


petrosal sinus 
enlarged vein that receives blood from the cavernous sinus and flows 
into the internal jugular vein 


phrenic vein 
drains the diaphragm; the right phrenic vein flows into the inferior 
vena cava and the left phrenic vein leads to the left renal vein 


plantar arch 
formed from the anastomosis of the dorsalis pedis artery and medial 
and plantar arteries; branches supply the distal portions of the foot and 
digits 


plantar veins 
drain the foot and lead to the plantar venous arch 


plantar venous arch 
formed from the plantar veins; leads to the anterior and posterior tibial 
veins through anastomoses 


popliteal artery 
continuation of the femoral artery posterior to the knee; branches into 
the anterior and posterior tibial arteries 


popliteal vein 
continuation of the femoral vein behind the knee; drains the region 
behind the knee and forms from the fusion of the fibular and anterior 
and posterior tibial veins 


posterior cerebral artery 
branch of the basilar artery that forms a portion of the posterior 
segment of the arterial circle; supplies blood to the posterior portion of 
the cerebrum and brain stem 


posterior communicating artery 


branch of the posterior cerebral artery that forms part of the posterior 
portion of the arterial circle; supplies blood to the brain 


posterior tibial artery 
branch from the popliteal artery that gives rise to the fibular or 
peroneal artery; supplies blood to the posterior tibial region 


posterior tibial vein 
forms from the dorsal venous arch; drains the area near the posterior 
surface of the tibia and leads to the popliteal vein 


pulmonary artery 
one of two branches, left and right, that divides off from the pulmonary 
trunk and leads to smaller arterioles and eventually to the pulmonary 
capillaries 


pulmonary circuit 
system of blood vessels that provide gas exchange via a network of 
arteries, veins, and capillaries that run from the heart, through the 
body, and back to the lungs 


pulmonary trunk 
single large vessel exiting the right ventricle that divides to form the 
right and left pulmonary arteries 


pulmonary veins 
two sets of paired vessels, one pair on each side, that are formed from 
the small venules leading away from the pulmonary capillaries that 
flow into the left atrium 


radial artery 
formed at the bifurcation of the brachial artery; parallels the radius; 
gives off smaller branches until it reaches the carpal region where it 
fuses with the ulnar artery to form the superficial and deep palmar 
arches; supplies blood to the lower arm and carpal region 


radial vein 


parallels the radius and radial artery; arises from the palmar venous 
arches and leads to the brachial vein 


renal artery 
branch of the abdominal aorta; supplies each kidney 


renal vein 
largest vein entering the inferior vena cava; drains the kidneys and 
leads to the inferior vena cava 


right gastric artery 
branch of the common hepatic artery; supplies blood to the stomach 


sigmoid sinuses 
enlarged veins that receive blood from the transverse sinuses; flow 
through the jugular foramen and into the internal jugular vein 


small saphenous vein 
located on the lateral surface of the leg; drains blood from the 
superficial regions of the lower leg and foot, and leads to the popliteal 
vein 


splenic artery 
branch of the celiac trunk; supplies blood to the spleen 


straight sinus 
enlarged vein that drains blood from the brain; receives most of the 
blood from the great cerebral vein and flows into the left or right 
transverse sinus 


subclavian artery 
right subclavian arises from the brachiocephalic artery, whereas the left 
subclavian artery arises from the aortic arch; gives rise to the internal 
thoracic, vertebral, and thyrocervical arteries; supplies blood to the 
arms, chest, shoulders, back, and central nervous system 


subclavian vein 


located deep in the thoracic cavity; becomes the axillary vein as it 
enters the axillary region; drains the axillary and smaller local veins 
near the scapular region; leads to the brachiocephalic vein 


subscapular vein 
drains blood from the subscapular region and leads to the axillary vein 


superior mesenteric artery 
branch of the abdominal aorta; supplies blood to the small intestine 
(duodenum, jejunum, and ileum), the pancreas, and a majority of the 
large intestine 


superior phrenic artery 
branch of the thoracic aorta; supplies blood to the superior surface of 
the diaphragm 


superior sagittal sinus 
enlarged vein located midsagittally between the meningeal and 
periosteal layers of the dura mater within the falx cerebri; receives 
most of the blood drained from the superior surface of the cerebrum 
and leads to the inferior jugular vein and the vertebral vein 


superior vena cava 
large systemic vein; drains blood from most areas superior to the 
diaphragm; empties into the right atrium 


temporal vein 
drains blood from the temporal region and leads to the external jugular 
vein 


testicular artery 
branch of the abdominal aorta; will ultimately travel outside the body 
cavity to the testes and form one component of the spermatic cord 


testicular vein 
drains the testes and forms part of the spermatic cord; the right 
testicular vein empties directly into the inferior vena cava and the left 
testicular vein empties into the left renal vein 


thoracic aorta 
portion of the descending aorta superior to the aortic hiatus 


thyrocervical artery 
arises from the subclavian artery; supplies blood to the thyroid, the 
cervical region, the upper back, and shoulder 


transient ischemic attack (TIA) 
temporary loss of neurological function caused by a brief interruption 
in blood flow; also known as a mini-stroke 


transverse sinuses 
pair of enlarged veins near the lambdoid suture that drain the occipital, 
Sagittal, and straight sinuses, and leads to the sigmoid sinuses 


trunk 
large vessel that gives rise to smaller vessels 


ulnar artery 
formed at the bifurcation of the brachial artery; parallels the ulna; 
gives off smaller branches until it reaches the carpal region where it 
fuses with the radial artery to form the superficial and deep palmar 
arches; supplies blood to the lower arm and carpal region 


ulnar vein 
parallels the ulna and ulnar artery; arises from the palmar venous 
arches and leads to the brachial vein 


vertebral artery 
arises from the subclavian artery and passes through the vertebral 
foramen through the foramen magnum to the brain; joins with the 
internal carotid artery to form the arterial circle; supplies blood to the 
brain and spinal cord 


vertebral vein 
arises from the base of the brain and the cervical region of the spinal 
cord; passes through the intervertebral foramina in the cervical 
vertebrae; drains smaller veins from the cranium, spinal cord, and 


vertebrae, and leads to the brachiocephalic vein; counterpart of the 
vertebral artery 


visceral branches 
branches of the descending aorta that supply blood to the viscera 


An Overview of Blood 
By the end of this section, you will be able to: 


e Identify the primary functions of blood in transportation, defense, and 
maintenance of homeostasis 

e Name the fluid component of blood and the three major types of 
formed elements, and identify their relative proportions in a blood 
sample 

e Discuss the unique physical characteristics of blood 

e Identify the composition of blood plasma, including its most important 
solutes and plasma proteins 


Recall that blood is a connective tissue. Like all connective tissues, it is 
made up of cellular elements and an extracellular matrix. The cellular 
elements—referred to as the formed elements—include red blood cells 
(RBCs), white blood cells (WBCs), and cell fragments called platelets. 
The extracellular matrix, called plasma, makes blood unique among 
connective tissues because it is fluid. This fluid, which is mostly water, 
perpetually suspends the formed elements and enables them to circulate 
throughout the body within the cardiovascular system. 


Functions of Blood 


The primary function of blood is to deliver oxygen and nutrients to and 
remove wastes from body cells, but that is only the beginning of the story. 
The specific functions of blood also include defense, distribution of heat, 
and maintenance of homeostasis. 


Transportation 


Nutrients from the foods you eat are absorbed in the digestive tract. Most of 
these travel in the bloodstream directly to the liver, where they are 
processed and released back into the bloodstream for delivery to body cells. 
Oxygen from the air you breathe diffuses into the blood, which moves from 
the lungs to the heart, which then pumps it out to the rest of the body. 
Moreover, endocrine glands scattered throughout the body release their 


products, called hormones, into the bloodstream, which carries them to 
distant target cells. Blood also picks up cellular wastes and byproducts, and 
transports them to various organs for removal. For instance, blood moves 
carbon dioxide to the lungs for exhalation from the body, and various waste 
products are transported to the kidneys and liver for excretion from the 
body in the form of urine or bile. 


Defense 


Many types of WBCs protect the body from external threats, such as 
disease-causing bacteria that have entered the bloodstream in a wound. 
Other WBCs seek out and destroy internal threats, such as cells with 
mutated DNA that could multiply to become cancerous, or body cells 
infected with viruses. 


When damage to the vessels results in bleeding, blood platelets and certain 
proteins dissolved in the plasma, the fluid portion of the blood, interact to 
block the ruptured areas of the blood vessels involved. This protects the 
body from further blood loss. 


Maintenance of Homeostasis 


Recall that body temperature is regulated via a classic negative-feedback 
loop. If you were exercising on a warm day, your rising core body 
temperature would trigger several homeostatic mechanisms, including 
increased transport of blood from your core to your body periphery, which 
is typically cooler. As blood passes through the vessels of the skin, heat 
would be dissipated to the environment, and the blood returning to your 
body core would be cooler. In contrast, on a cold day, blood is diverted 
away from the skin to maintain a warmer body core. In extreme cases, this 
may result in frostbite. 


Blood also helps to maintain the chemical balance of the body. Proteins and 
other compounds in blood act as buffers, which thereby help to regulate the 


pH of body tissues. Blood also helps to regulate the water content of body 
cells. 


Composition of Blood 


You have probably had blood drawn from a superficial vein in your arm, 
which was then sent to a lab for analysis. Some of the most common blood 
tests—for instance, those measuring lipid or glucose levels in plasma— 
determine which substances are present within blood and in what quantities. 
Other blood tests check for the composition of the blood itself, including 
the quantities and types of formed elements. 


One such test, called a hematocrit, measures the percentage of RBCs, 
clinically known as erythrocytes, in a blood sample. It is performed by 
spinning the blood sample in a specialized centrifuge, a process that causes 
the heavier elements suspended within the blood sample to separate from 
the lightweight, liquid plasma ({link]). Because the heaviest elements in 
blood are the erythrocytes, these settle at the very bottom of the hematocrit 
tube. Located above the erythrocytes is a pale, thin layer composed of the 
remaining formed elements of blood. These are the WBCGs, clinically 
known as leukocytes, and the platelets, cell fragments also called 
thrombocytes. This layer is referred to as the buffy coat because of its 
color; it normally constitutes less than 1 percent of a blood sample. Above 
the buffy coat is the blood plasma, normally a pale, straw-colored fluid, 
which constitutes the remainder of the sample. 


The volume of erythrocytes after centrifugation is also commonly referred 
to as packed cell volume (PCV). In normal blood, about 45 percent of a 
sample is erythrocytes. The hematocrit of any one sample can vary 
significantly, however, about 36—50 percent, according to gender and other 
factors. Normal hematocrit values for females range from 37 to 47, with a 
mean value of 41; for males, hematocrit ranges from 42 to 52, with a mean 
of 47. The percentage of other formed elements, the WBCs and platelets, is 
extremely small so it is not normally considered with the hematocrit. So the 
mean plasma percentage is the percent of blood that is not erythrocytes: for 
females, it is approximately 59 (or 100 minus 41), and for males, it is 
approximately 53 (or 100 minus 47). 


Composition of Blood 


Plasma: 
- Water, proteins, 
nutrients, hormones, 
etc. 


Buffy coat: 
- White blood cells, 
platelets 


Hematocrit: 
- Red blood cells 


Normal Blood: Anemia: Polycythemia: 
Q. 37%-47% hematocrit Depressed Elevated 
O" 42%-52% hematocrit hematocrit % hematocrit % 


The cellular elements of blood include a 
vast number of erythrocytes and 
comparatively fewer leukocytes and 
platelets. Plasma is the fluid in which the 
formed elements are suspended. A sample 
of blood spun in a centrifuge reveals that 
plasma is the lightest component. It floats 
at the top of the tube separated from the 
heaviest elements, the erythrocytes, by a 
buffy coat of leukocytes and platelets. 
Hematocrit is the percentage of the total 
sample that is comprised of erythrocytes. 
Depressed and elevated hematocrit levels 
are shown for comparison. 


Characteristics of Blood 


When you think about blood, the first characteristic that probably comes to 
mind is its color. Blood that has just taken up oxygen in the lungs is bright 
red, and blood that has released oxygen in the tissues is a more dusky red. 
This is because hemoglobin is a pigment that changes color, depending 
upon the degree of oxygen saturation. 


Blood is viscous and somewhat sticky to the touch. It has a viscosity 
approximately five times greater than water. Viscosity is a measure of a 
fluid’s thickness or resistance to flow, and is influenced by the presence of 
the plasma proteins and formed elements within the blood. The viscosity of 
blood has a dramatic impact on blood pressure and flow. Consider the 
difference in flow between water and honey. The more viscous honey 
would demonstrate a greater resistance to flow than the less viscous water. 
The same principle applies to blood. 


The normal temperature of blood is slightly higher than normal body 
temperature—about 38 °C (or 100.4 °F), compared to 37 °C (or 98.6 °F) for 
an internal body temperature reading, although daily variations of 0.5 °C 
are normal. Although the surface of blood vessels is relatively smooth, as 
blood flows through them, it experiences some friction and resistance, 
especially as vessels age and lose their elasticity, thereby producing heat. 
This accounts for its slightly higher temperature. 


The pH of blood averages about 7.4; however, it can range from 7.35 to 
7.45 in a healthy person. Blood is therefore somewhat more basic (alkaline) 
on a chemical scale than pure water, which has a pH of 7.0. Blood contains 
numerous buffers that actually help to regulate pH. 


Blood constitutes approximately 8 percent of adult body weight. Adult 
males typically average about 5 to 6 liters of blood. Females average 4—5 
liters. 


Blood Plasma 


Like other fluids in the body, plasma is composed primarily of water: In 
fact, it is about 92 percent water. Dissolved or suspended within this water 
is a mixture of substances, most of which are proteins. There are literally 
hundreds of substances dissolved or suspended in the plasma, although 
many of them are found only in very small quantities. 


Note: 


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inten 

Visit this site for a list of normal levels established for many of the 
substances found in a sample of blood. Serum, one of the specimen types 
included, refers to a sample of plasma after clotting factors have been 
removed. What types of measurements are given for levels of glucose in 
the blood? 


Plasma Proteins 


About 7 percent of the volume of plasma—nearly all that is not water—is 
made of proteins. These include several plasma proteins (proteins that are 
unique to the plasma), plus a much smaller number of regulatory proteins, 
including enzymes and some hormones. The major components of plasma 
are summarized in [link]. 


The three major groups of plasma proteins are as follows: 


e Albumin is the most abundant of the plasma proteins. Manufactured 
by the liver, albumin molecules serve as binding proteins—transport 
vehicles for fatty acids and steroid hormones. Recall that lipids are 
hydrophobic; however, their binding to albumin enables their transport 
in the watery plasma. Albumin is also the most significant contributor 
to the osmotic pressure of blood; that is, its presence holds water inside 
the blood vessels and draws water from the tissues, across blood vessel 
walls, and into the bloodstream. This in turn helps to maintain both 
blood volume and blood pressure. Albumin normally accounts for 
approximately 54 percent of the total plasma protein content, in 
clinical levels of 3.5-5.0 g/dL blood. 

e The second most common plasma proteins are the globulins. A 
heterogeneous group, there are three main subgroups known as alpha, 
beta, and gamma globulins. The alpha and beta globulins transport 


iron, lipids, and the fat-soluble vitamins A, D, E, and K to the cells; 
like albumin, they also contribute to osmotic pressure. The gamma 
globulins are proteins involved in immunity and are better known as an 
antibodies or immunoglobulins. Although other plasma proteins are 
produced by the liver, immunoglobulins are produced by specialized 
leukocytes known as plasma cells. (Seek additional content for more 
information about immunoglobulins.) Globulins make up 
approximately 38 percent of the total plasma protein volume, in 
clinical levels of 1.0—1.5 g/dL blood. 

e The least abundant plasma protein is fibrinogen. Like albumin and the 
alpha and beta globulins, fibrinogen is produced by the liver. It is 
essential for blood clotting, a process described later in this chapter. 
Fibrinogen accounts for about 7 percent of the total plasma protein 
volume, in clinical levels of 0.2—-0.45 g/dL blood. 


Other Plasma Solutes 


In addition to proteins, plasma contains a wide variety of other substances. 
These include various electrolytes, such as sodium, potassium, and calcium 
ions; dissolved gases, such as oxygen, carbon dioxide, and nitrogen; various 
organic nutrients, such as vitamins, lipids, glucose, and amino acids; and 
metabolic wastes. All of these nonprotein solutes combined contribute 
approximately 1 percent to the total volume of plasma. 

Major Blood Components 


Component Subcomponent Type and % 
and % and % of (where Site of production Major function(s) 
of blood component appropriate) 


Absorbed by intestinal 
tract or produced by 
metabolism 


Water 
92 percent 


Transport 
medium 


Maintain osmotic 
Albumin concentration, 
54-60 percent transport lipid 
molecules 


Transport, 
maintain osmotic 
concentration 


Alpha globulins— 
liver 


Plasma proteins 
Plasma 7 percent Globulins Beta globulins— 
46-63 35-38 percent liver 
percent 


Transport, 
maintain osmotic 
concentration 


Gamma globulins 
(immunoglobulins) 
—plasma cells 


Immune 
responses 


Fibrinogen Blood clotting in 
4-7 percent hemostasis 


Regulatory proteins Hormones : Regulate various 
and enzymes Various sources body functions 


Absorbed by intestinal 
Other solutes Nutrients, gases, tract, exchanged in Numerous 
1 percent and wastes respiratory system, and varied 
or produced by cells 


Transport gases, 

Erythrocytes primarily oxygen 

99 percent Erytwocyive Red bone marrow and some carbon 
dioxide 


Granular 
leukocytes: 

neutrophils Red bone marrow 
eosinophils 

basophils 


Nonspecific 
Formed immunity 
elements Leukocytes 
37-54 <1 percent 
percent Platelets Lymphocytes: Lymphocytes: 
<1 percent Agranular bone marrow and specific 
leukocytes: lymphatic tissue immunity 
lymphocytes 
monocytes Monocytes: Monocytes: 
red bone marrow nonspecific immunity 


Platelets Megakaryocytes: : 


Note: 

Career Connection 

Phlebotomy and Medical Lab Technology 

Phlebotomists are professionals trained to draw blood (phleb- = “a blood 
vessel”; -tomy = “to cut”). When more than a few drops of blood are 


required, phlebotomists perform a venipuncture, typically of a surface vein 
in the arm. They perform a capillary stick on a finger, an earlobe, or the 
heel of an infant when only a small quantity of blood is required. An 
arterial stick is collected from an artery and used to analyze blood gases. 
After collection, the blood may be analyzed by medical laboratories or 
perhaps used for transfusions, donations, or research. While many allied 
health professionals practice phlebotomy, the American Society of 
Phlebotomy Technicians issues certificates to individuals passing a 
national examination, and some large labs and hospitals hire individuals 
expressly for their skill in phlebotomy. 

Medical or clinical laboratories employ a variety of individuals in technical 
positions: 


e Medical technologists (MT), also known as clinical laboratory 
technologists (CLT), typically hold a bachelor’s degree and 
certification from an accredited training program. They perform a 
wide variety of tests on various body fluids, including blood. The 
information they provide is essential to the primary care providers in 
determining a diagnosis and in monitoring the course of a disease and 
response to treatment. 

¢ Medical laboratory technicians (MLT) typically have an associate’s 
degree but may perform duties similar to those of an MT. 

e Medical laboratory assistants (MLA) spend the majority of their time 
processing samples and carrying out routine assignments within the 
lab. Clinical training is required, but a degree may not be essential to 
obtaining a position. 


Chapter Review 


Blood is a fluid connective tissue critical to the transportation of nutrients, 
gases, and wastes throughout the body; to defend the body against infection 
and other threats; and to the homeostatic regulation of pH, temperature, and 
other internal conditions. Blood is composed of formed elements— 
erythrocytes, leukocytes, and cell fragments called platelets—and a fluid 
extracellular matrix called plasma. More than 90 percent of plasma is water. 


The remainder is mostly plasma proteins—mainly albumin, globulins, and 
fibrinogen—and other dissolved solutes such as glucose, lipids, electrolytes, 
and dissolved gases. Because of the formed elements and the plasma 
proteins and other solutes, blood is sticky and more viscous than water. It is 
also slightly alkaline, and its temperature is slightly higher than normal 
body temperature. 


Interactive Link Questions 


Exercise: 


Problem: 


Visit this site for a list of normal levels established for many of the 
substances found in a sample of blood. Serum, one of the specimen 
types included, refers to a sample of plasma after clotting factors have 
been removed. What types of measurements are given for levels of 
glucose in the blood? 


Solution: 
There are values given for percent saturation, tension, and blood gas, 
and there are listings for different types of hemoglobin. 

Review Questions 


Exercise: 


Problem: Which of the following statements about blood is true? 


a. Blood is about 92 percent water. 

b. Blood is slightly more acidic than water. 
c. Blood is slightly more viscous than water. 
d. Blood is slightly more salty than seawater. 


Solution: 


C 
Exercise: 
Problem: Which of the following statements about albumin is true? 


a. It draws water out of the blood vessels and into the body’s tissues. 
b. It is the most abundant plasma protein. 

c. It is produced by specialized leukocytes called plasma cells. 

d. All of the above are true. 


Solution: 


B 
Exercise: 


Problem: 
Which of the following plasma proteins is not produced by the liver? 
a. fibrinogen 
b. alpha globulin 
c. beta globulin 
d. immunoglobulin 


Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: 


A patient’s hematocrit is 42 percent. Approximately what percentage 
of the patient’s blood is plasma? 


Solution: 


The patient’s blood is approximately 58 percent plasma (since the 
buffy coat is less than 1 percent). 
Exercise: 


Problem: 

Why would it be incorrect to refer to the formed elements as cells? 
Solution: 

The formed elements include erythrocytes and leukocytes, which are 
cells (although mature erythrocytes do not have a nucleus); however, 


the formed elements also include platelets, which are not true cells but 
cell fragments. 


Exercise: 


Problem: 


True or false: The buffy coat is the portion of a blood sample that is 
made up of its proteins. 


Solution: 


False. The buffy coat is the portion of blood that is made up of its 
leukocytes and platelets. 


Glossary 


albumin 


most abundant plasma protein, accounting for most of the osmotic 
pressure of plasma 


antibodies 
(also, immunoglobulins or gamma globulins) antigen-specific proteins 
produced by specialized B lymphocytes that protect the body by 
binding to foreign objects such as bacteria and viruses 


blood 
liquid connective tissue composed of formed elements—erythrocytes, 
leukocytes, and platelets—and a fluid extracellular matrix called 
plasma; component of the cardiovascular system 


buffy coat 
thin, pale layer of leukocytes and platelets that separates the 
erythrocytes from the plasma in a sample of centrifuged blood 


fibrinogen 
plasma protein produced in the liver and involved in blood clotting 


formed elements 
cellular components of blood; that is, erythrocytes, leukocytes, and 
platelets 


globulins 
heterogeneous group of plasma proteins that includes transport 
proteins, clotting factors, immune proteins, and others 


hematocrit 
(also, packed cell volume) volume percentage of erythrocytes in a 
sample of centrifuged blood 


immunoglobulins 
(also, antibodies or gamma globulins) antigen-specific proteins 
produced by specialized B lymphocytes that protect the body by 
binding to foreign objects such as bacteria and viruses 


packed cell volume (PCV) 


(also, hematocrit) volume percentage of erythrocytes present in a 
sample of centrifuged blood 


plasma 
in blood, the liquid extracellular matrix composed mostly of water that 
circulates the formed elements and dissolved materials throughout the 
cardiovascular system 


platelets 
(also, thrombocytes) one of the formed elements of blood that consists 
of cell fragments broken off from megakaryocytes 


red blood cells (RBCs) 
(also, erythrocytes) one of the formed elements of blood that transports 
oxygen 


white blood cells (WBCs) 
(also, leukocytes) one of the formed elements of blood that provides 
defense against disease agents and foreign materials 


Production of the Formed Elements 
By the end of this section, you will be able to: 


e Trace the generation of the formed elements of blood from bone 
marrow stem cells 

e Discuss the role of hemopoietic growth factors in promoting the 
production of the formed elements 


The lifespan of the formed elements is very brief. Although one type of 
leukocyte called memory cells can survive for years, most erythrocytes, 
leukocytes, and platelets normally live only a few hours to a few weeks. 
Thus, the body must form new blood cells and platelets quickly and 
continuously. When you donate a unit of blood during a blood drive 
(approximately 475 mL, or about 1 pint), your body typically replaces the 
donated plasma within 24 hours, but it takes about 4 to 6 weeks to replace 
the blood cells. This restricts the frequency with which donors can 
contribute their blood. The process by which this replacement occurs is 
called hemopoiesis, or hematopoiesis (from the Greek root haima- = 
“blood”; -poiesis = “production”). 


Sites of Hemopoiesis 


Prior to birth, hemopoiesis occurs in a number of tissues, beginning with 
the yolk sac of the developing embryo, and continuing in the fetal liver, 
spleen, lymphatic tissue, and eventually the red bone marrow. Following 
birth, most hemopoiesis occurs in the red marrow, a connective tissue 
within the spaces of spongy (cancellous) bone tissue. In children, 
hemopoiesis can occur in the medullary cavity of long bones; in adults, the 
process is largely restricted to the cranial and pelvic bones, the vertebrae, 
the sternum, and the proximal epiphyses of the femur and humerus. 


Throughout adulthood, the liver and spleen maintain their ability to 
generate the formed elements. This process is referred to as extramedullary 
hemopoiesis (meaning hemopoiesis outside the medullary cavity of adult 
bones). When a disease such as bone cancer destroys the bone marrow, 
causing hemopoiesis to fail, extramedullary hemopoiesis may be initiated. 


Differentiation of Formed Elements from Stem Cells 


All formed elements arise from stem cells of the red bone marrow. Recall 
that stem cells undergo mitosis plus cytokinesis (cellular division) to give 
rise to new daughter cells: One of these remains a stem cell and the other 
differentiates into one of any number of diverse cell types. Stem cells may 
be viewed as occupying a hierarchal system, with some loss of the ability to 
diversify at each step. The totipotent stem cell is the zygote, or fertilized 
egg. The totipotent (toti- = “all”) stem cell gives rise to all cells of the 
human body. The next level is the pluripotent stem cell, which gives rise 
to multiple types of cells of the body and some of the supporting fetal 
membranes. Beneath this level, the mesenchymal cell is a stem cell that 
develops only into types of connective tissue, including fibrous connective 
tissue, bone, cartilage, and blood, but not epithelium, muscle, and nervous 
tissue. One step lower on the hierarchy of stem cells is the hemopoietic 
stem cell, or hemocytoblast. All of the formed elements of blood originate 
from this specific type of cell. 


Hemopoiesis begins when the hemopoietic stem cell is exposed to 
appropriate chemical stimuli collectively called hemopoietic growth 
factors, which prompt it to divide and differentiate. One daughter cell 
remains a hemopoietic stem cell, allowing hemopoiesis to continue. The 
other daughter cell becomes either of two types of more specialized stem 
cells ([link]): 


¢ Lymphoid stem cells give rise to a class of leukocytes known as 
lymphocytes, which include the various T cells, B cells, and natural 
killer (NK) cells, all of which function in immunity. However, 
hemopoiesis of lymphocytes progresses somewhat differently from the 
process for the other formed elements. In brief, lymphoid stem cells 
quickly migrate from the bone marrow to lymphatic tissues, including 
the lymph nodes, spleen, and thymus, where their production and 
differentiation continues. B cells are so named since they mature in the 
bone marrow, while T cells mature in the thymus. 

¢ Myeloid stem cells give rise to all the other formed elements, 
including the erythrocytes; megakaryocytes that produce platelets; and 


a myeloblast lineage that gives rise to monocytes and three forms of 
granular leukocytes: neutrophils, eosinophils, and basophils. 


Hematopoietic System of Bone Marrow 


@ 
Multipotent hematopoietic 
stem cell (hemocytoblast) 


After division, some cells 
remain stem cells. 


@ i Ng The remaining cell goes down one of two paths 


depending on the chemical signals received. 


OF 


Myeloid stem cell Lymphoid stem cell 


rs rs 
o ~) @ @ ‘ 


isn a a a a 


Megakaryocyte Erythrocyte Basophil Neutrophil Eosinophil Monocyte Q. 
T lymphocyte —_B lymphocyte 


Megakaryoblast Proerythroblast Myeloblast Monoblast Lymphoblast 
Reticulocyte = ak 
rad | & 
i_- Natural killer cell Small lymphocyte 
~ (large granular 


xy’ 
Sy 
wp 
FN 


Platelets 


Hemopoiesis is the proliferation and differentiation of the formed 
elements of blood. 


Lymphoid and myeloid stem cells do not immediately divide and 
differentiate into mature formed elements. As you can see in [link], there 
are several intermediate stages of precursor cells (literally, forerunner cells), 
many of which can be recognized by their names, which have the suffix - 
blast. For instance, megakaryoblasts are the precursors of megakaryocytes, 
and proerythroblasts become reticulocytes, which eject their nucleus and 
most other organelles before maturing into erythrocytes. 


Hemopoietic Growth Factors 


Development from stem cells to precursor cells to mature cells is again 
initiated by hemopoietic growth factors. These include the following: 


e Erythropoietin (EPO) is a glycoprotein hormone secreted by the 
interstitial fibroblast cells of the kidneys in response to low oxygen 
levels. It prompts the production of erythrocytes. Some athletes use 
synthetic EPO as a performance-enhancing drug (called blood doping) 
to increase RBC counts and subsequently increase oxygen delivery to 
tissues throughout the body. EPO is a banned substance in most 
organized sports, but it is also used medically in the treatment of 
certain anemia, specifically those triggered by certain types of cancer, 
and other disorders in which increased erythrocyte counts and oxygen 
levels are desirable. 

e¢ Thrombopoietin, another glycoprotein hormone, is produced by the 
liver and kidneys. It triggers the development of megakaryocytes into 
platelets. 

¢ Cytokines are glycoproteins secreted by a wide variety of cells, 
including red bone marrow, leukocytes, macrophages, fibroblasts, and 
endothelial cells. They act locally as autocrine or paracrine factors, 
stimulating the proliferation of progenitor cells and helping to 
stimulate both nonspecific and specific resistance to disease. There are 
two major subtypes of cytokines known as colony-stimulating factors 
and interleukins. 


o Colony-stimulating factors (CSFs) are glycoproteins that act 
locally, as autocrine or paracrine factors. Some trigger the 
differentiation of myeloblasts into granular leukocytes, namely, 
neutrophils, eosinophils, and basophils. These are referred to as 
granulocyte CSFs. A different CSF induces the production of 
monocytes, called monocyte CSFs. Both granulocytes and 
monocytes are stimulated by GM-CSF; granulocytes, monocytes, 
platelets, and erythrocytes are stimulated by multi-CSF. Synthetic 
forms of these hormones are often administered to patients with 
various forms of cancer who are receiving chemotherapy to 
revive their WBC counts. 


o Interleukins are another class of cytokine signaling molecules 
important in hemopoiesis. They were initially thought to be 
secreted uniquely by leukocytes and to communicate only with 
other leukocytes, and were named accordingly, but are now 
known to be produced by a variety of cells including bone 
marrow and endothelium. Researchers now suspect that 
interleukins may play other roles in body functioning, including 
differentiation and maturation of cells, producing immunity and 
inflammation. To date, more than a dozen interleukins have been 
identified, with others likely to follow. They are generally 
numbered IL-1, IL-2, IL-3, etc. 


Note: 

Everyday Connection 

Blood Doping 

In its original intent, the term blood doping was used to describe the 
practice of injecting by transfusion supplemental RBCs into an individual, 
typically to enhance performance in a sport. Additional RBCs would 
deliver more oxygen to the tissues, providing extra aerobic capacity, 
clinically referred to as VO» max. The source of the cells was either from 
the recipient (autologous) or from a donor with compatible blood 
(homologous). This practice was aided by the well-developed techniques 
of harvesting, concentrating, and freezing of the RBCs that could be later 
thawed and injected, yet still retain their functionality. These practices are 
considered illegal in virtually all sports and run the risk of infection, 
significantly increasing the viscosity of the blood and the potential for 
transmission of blood-bome pathogens if the blood was collected from 
another individual. 

With the development of synthetic EPO in the 1980s, it became possible to 
provide additional RBCs by artificially stimulating RBC production in the 
bone marrow. Originally developed to treat patients suffering from anemia, 
renal failure, or cancer treatment, large quantities of EPO can be generated 
by recombinant DNA technology. Synthetic EPO is injected under the skin 
and can increase hematocrit for many weeks. It may also induce 
polycythemia and raise hematocrit to 70 or greater. This increased 


viscosity raises the resistance of the blood and forces the heart to pump 
more powerfully; in extreme cases, it has resulted in death. Other drugs 
such as cobalt II chloride have been shown to increase natural EPO gene 
expression. Blood doping has become problematic in many sports, 
especially cycling. Lance Armstrong, winner of seven Tour de France and 
many other cycling titles, was stripped of his victories and admitted to 
blood doping in 2013. 


Note: 


we 


— 
meee OPENStAX COLLEGE 


Watch this video to see doctors discuss the dangers of blood doping in 
sports. What are the some potential side effects of blood doping? 


Bone Marrow Sampling and Transplants 


Sometimes, a healthcare provider will order a bone marrow biopsy, a 
diagnostic test of a sample of red bone marrow, or a bone marrow 
transplant, a treatment in which a donor’s healthy bone marrow—and its 
stem cells—replaces the faulty bone marrow of a patient. These tests and 
procedures are often used to assist in the diagnosis and treatment of various 
severe forms of anemia, such as thalassemia major and sickle cell anemia, 
as well as some types of cancer, specifically leukemia. 


In the past, when a bone marrow sample or transplant was necessary, the 
procedure would have required inserting a large-bore needle into the region 
near the iliac crest of the pelvic bones (os coxae). This location was 
preferred, since its location close to the body surface makes it more 


accessible, and it is relatively isolated from most vital organs. 
Unfortunately, the procedure is quite painful. 


Now, direct sampling of bone marrow can often be avoided. In many cases, 
stem cells can be isolated in just a few hours from a sample of a patient’s 
blood. The isolated stem cells are then grown in culture using the 
appropriate hemopoietic growth factors, and analyzed or sometimes frozen 
for later use. 


For an individual requiring a transplant, a matching donor is essential to 
prevent the immune system from destroying the donor cells—a 
phenomenon known as tissue rejection. To treat patients with bone marrow 
transplants, it is first necessary to destroy the patient’s own diseased 
marrow through radiation and/or chemotherapy. Donor bone marrow stem 
cells are then intravenously infused. From the bloodstream, they establish 
themselves in the recipient’s bone marrow. 


Chapter Review 


Through the process of hemopoiesis, the formed elements of blood are 
continually produced, replacing the relatively short-lived erythrocytes, 
leukocytes, and platelets. Hemopoiesis begins in the red bone marrow, with 
hemopoietic stem cells that differentiate into myeloid and lymphoid 
lineages. Myeloid stem cells give rise to most of the formed elements. 
Lymphoid stem cells give rise only to the various lymphocytes designated 
as B and T cells, and NK cells. Hemopoietic growth factors, including 
erythropoietin, thrombopoietin, colony-stimulating factors, and interleukins, 
promote the proliferation and differentiation of formed elements. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to see doctors discuss the dangers of blood doping in 
sports. What are the some potential side effects of blood doping? 


Solution: 


Side effects can include heart disease, stroke, pulmonary embolism, 
and virus transmission. 


Review Questions 


Exercise: 


Problem: 
Which of the formed elements arise from myeloid stem cells? 


a. B cells 

b. natural killer cells 
c. platelets 

d. all of the above 


Solution: 


C 
Exercise: 


Problem: 
Which of the following statements about erythropoietin is true? 


a. It facilitates the proliferation and differentiation of the erythrocyte 
lineage. 

b. It is a hormone produced by the thyroid gland. 

c. It is a hemopoietic growth factor that prompts lymphoid stem 
cells to leave the bone marrow. 

d. Both a and b are true. 


Solution: 


A 
Exercise: 


Problem: 
Interleukins are associated primarily with which of the following? 


a. production of various lymphocytes 
b. immune responses 

c. inflammation 

d. all of the above 


Solution: 


D 


Critical Thinking Questions 


Exercise: 
Problem: 
Myelofibrosis is a disorder in which inflammation and scar tissue 


formation in the bone marrow impair hemopoiesis. One sign is an 
enlarged spleen. Why? 


Solution: 


When disease impairs the ability of the bone marrow to participate in 
hemopoiesis, extramedullary hemopoiesis begins in the patient’s liver 
and spleen. This causes the spleen to enlarge. 


Exercise: 


Problem: 


Would you expect a patient with a form of cancer called acute 
myelogenous leukemia to experience impaired production of 
erythrocytes, or impaired production of lymphocytes? Explain your 
choice. 


Solution: 


The adjective myelogenous suggests a condition originating from 
(generated by) myeloid cells. Acute myelogenous leukemia impairs the 
production of erythrocytes and other mature formed elements of the 
myeloid stem cell lineage. Lymphocytes arise from the lymphoid stem 
cell line. 


Glossary 


bone marrow biopsy 
diagnostic test of a sample of red bone marrow 


bone marrow transplant 
treatment in which a donor’s healthy bone marrow with its stem cells 
replaces diseased or damaged bone marrow of a patient 


colony-stimulating factors (CSFs) 
glycoproteins that trigger the proliferation and differentiation of 
myeloblasts into granular leukocytes (basophils, neutrophils, and 
eosinophils) 


cytokines 
class of proteins that act as autocrine or paracrine signaling molecules; 
in the cardiovascular system, they stimulate the proliferation of 
progenitor cells and help to stimulate both nonspecific and specific 
resistance to disease 


erythropoietin (EPO) 


glycoprotein that triggers the bone marrow to produce RBCs; secreted 
by the kidney in response to low oxygen levels 


hemocytoblast 
hemopoietic stem cell that gives rise to the formed elements of blood 


hemopoiesis 
production of the formed elements of blood 


hemopoietic growth factors 
chemical signals including erythropoietin, thrombopoietin, colony- 
stimulating factors, and interleukins that regulate the differentiation 
and proliferation of particular blood progenitor cells 


hemopoietic stem cell 
type of pluripotent stem cell that gives rise to the formed elements of 
blood (hemocytoblast) 


interleukins 
signaling molecules that may function in hemopoiesis, inflammation, 
and specific immune responses 


lymphoid stem cells 
type of hemopoietic stem cells that gives rise to lymphocytes, 
including various T cells, B cells, and NK cells, all of which function 
in immunity 


myeloid stem cells 
type of hemopoietic stem cell that gives rise to some formed elements, 
including erythrocytes, megakaryocytes that produce platelets, and a 
myeloblast lineage that gives rise to monocytes and three forms of 
granular leukocytes (neutrophils, eosinophils, and basophils) 


pluripotent stem cell 
stem cell that derives from totipotent stem cells and is capable of 


differentiating into many, but not all, cell types 


totipotent stem cell 


embryonic stem cell that is capable of differentiating into any and all 
cells of the body; enabling the full development of an organism 


thrombopoietin 
hormone secreted by the liver and kidneys that prompts the 
development of megakaryocytes into thrombocytes (platelets) 


Erythrocytes 
By the end of this section, you will be able to: 


e Describe the anatomy of erythrocytes 
e Discuss the various steps in the lifecycle of an erythrocyte 
e Explain the composition and function of hemoglobin 


The erythrocyte, commonly known as a red blood cell (or RBC), is by far 
the most common formed element: A single drop of blood contains millions 
of erythrocytes and just thousands of leukocytes. Specifically, males have 
about 5.4 million erythrocytes per microliter (uL) of blood, and females 
have approximately 4.8 million per pL. In fact, erythrocytes are estimated 
to make up about 25 percent of the total cells in the body. As you can 
imagine, they are quite small cells, with a mean diameter of only about 7-8 
micrometers (um) ({link]). The primary functions of erythrocytes are to 
pick up inhaled oxygen from the lungs and transport it to the body’s tissues, 
and to pick up some (about 24 percent) carbon dioxide waste at the tissues 
and transport it to the lungs for exhalation. Erythrocytes remain within the 
vascular network. Although leukocytes typically leave the blood vessels to 
perform their defensive functions, movement of erythrocytes from the 
blood vessels is abnormal. 

Summary of Formed Elements in Blood 


Formed 
element 


Numbers Comments 
present per 
microliter (uL) 


and mean (range) 


Major 
subtypes 


Appearance ina 
standard 
blood smear 


Summary of 
functions 


Erythrocytes 
(red blood 


cells) . 


Flattened biconcave 
disk; no nucleus; 
pale red color 


5.2 million 
(4.4-6.0 million) 


Transport oxygen and 
some carbon dioxide 
between tissues and 
lungs 


Leukocytes 
(white blood 
cells) 


7000 
(5000—10,000) 


Obvious dark-staining 
nucleus 


All function in body 
defenses 


Exit capillaries and 
move into tissues; 
lifespan of usually 
a few hours or days 


4360 
(1800-9950) 


Granulocytes 
including 
neutrophils, 
eosinophils, and 
basophils 


Abundant granules in 
cytoplasm; nucleus 
normally lobed 


Nonspecific (innate) 
resistance to disease 


Classified according 
to membrane-bound 
granules in cytoplasm 


Neutrophils 


4150 
(1800-7300) 


Nuclear lobes 
increase with age; 
pale lilac granules 


Phagocytic; 
particularly effective 
against bacteria. 
Release cytotoxic 
chemicals from 


Most common 
leukocyte; 

lifespan of minutes 
to days 


granules 
Eosinophils 165 Nucleus generally Phagocytic cells; Lifespan of 
(0-700) two-lobed; bright particularly effective minutes to days 
La red-orange granules with antigen- antibody 


2640 
(1700-4950) 


2185 
(1500-4000) 


455 
(200-950) 


Nucleus generally 
two-lobed but difficult 
to see due to 
presence of heavy, 
dense, dark purple 
granules 


Lack abundant 
granules in cytoplasm; 
have a simple- 
shaped nucleus that 
may be indented 


Spherical cells with 

a single often large 
nucleus occupying 
much of the cell’s 
volume; stains purple; 
seen in large 

(natural killer cells) and 
small (B and T cells) 
variants 


Largest leukocyte 
with an indented or 
horseshoe-shaped 
nucleus 


complexes. Release 
antihistamines. 
Increase in allergies 
and parasitic infections 


Promotes 
inflammation 


Body defenses 


Primarily specific 
(adaptive) immunity: 
T cells directly attack 
other cells (cellular 
immunity); B cells 
release antibodies 
(humoral immunity); 
natural killer cells are 
similar to T cells but 
nonspecific 


Very effective 
phagocytic cells 
engulfing pathogens 
or worn out cells; also 


Least common 
leukocyte; lifespan 
unknown 


Group consists of 
two major cell types 
from different 
lineages 


Initial cells originate 

in bone marrow, but 
secondary production 
occurs in lymphatic 
tissue; several distinct 
subtypes; memory cells 
form after exposure to 
a pathogen and rapidly 
increase responses to 
subsequent exposure; 
lifespan of many years 


Produced in red bone 
marrow; referred to as 
macrophages after 
leaving circulation 


serve as antigen- 
presenting cells (APCs) 
for other components 
of the immune system 


350,000 Cellular fragments 

(150,000—500,000) | surrounded by a 
plasma membrane 
and containing 
granules; purple stain 


Hemostasis plus Formed from 

release growth factors }|megakaryocytes 

for repair and healing |that remain in the red 

of tissue bone marrow and shed 
platelets into circulation 


Platelets 


Shape and Structure of Erythrocytes 


As an erythrocyte matures in the red bone marrow, it extrudes its nucleus 
and most of its other organelles. During the first day or two that it is in the 
circulation, an immature erythrocyte, known as a reticulocyte, will still 
typically contain remnants of organelles. Reticulocytes should comprise 
approximately 1—2 percent of the erythrocyte count and provide a rough 
estimate of the rate of RBC production, with abnormally low or high rates 
indicating deviations in the production of these cells. These remnants, 
primarily of networks (reticulum) of ribosomes, are quickly shed, however, 
and mature, circulating erythrocytes have few internal cellular structural 
components. Lacking mitochondria, for example, they rely on anaerobic 
respiration. This means that they do not utilize any of the oxygen they are 
transporting, so they can deliver it all to the tissues. They also lack 


endoplasmic reticula and do not synthesize proteins. Erythrocytes do, 
however, contain some structural proteins that help the blood cells maintain 
their unique structure and enable them to change their shape to squeeze 
through capillaries. This includes the protein spectrin, a cytoskeletal protein 
element. 


Erythrocytes are biconcave disks; that is, they are plump at their periphery 
and very thin in the center ({link]). Since they lack most organelles, there is 
more interior space for the presence of the hemoglobin molecules that, as 
you will see shortly, transport gases. The biconcave shape also provides a 
greater surface area across which gas exchange can occur, relative to its 
volume; a sphere of a similar diameter would have a lower surface area-to- 
volume ratio. In the capillaries, the oxygen carried by the erythrocytes can 
diffuse into the plasma and then through the capillary walls to reach the 
cells, whereas some of the carbon dioxide produced by the cells as a waste 
product diffuses into the capillaries to be picked up by the erythrocytes. 
Capillary beds are extremely narrow, slowing the passage of the 
erythrocytes and providing an extended opportunity for gas exchange to 
occur. However, the space within capillaries can be so minute that, despite 
their own small size, erythrocytes may have to fold in on themselves if they 
are to make their way through. Fortunately, their structural proteins like 
spectrin are flexible, allowing them to bend over themselves to a surprising 
degree, then spring back again when they enter a wider vessel. In wider 
vessels, erythrocytes may stack up much like a roll of coins, forming a 
rouleaux, from the French word for “roll.” 

Shape of Red Blood Cells 


Erythrocytes are biconcave 
discs with very shallow centers. 
This shape optimizes the ratio 
of surface area to volume, 
facilitating gas exchange. It also 
enables them to fold up as they 
move through narrow blood 
vessels. 


Hemoglobin 


Hemoglobin is a large molecule made up of proteins and iron. It consists of 
four folded chains of a protein called globin, designated alpha 1 and 2, and 
beta 1 and 2 ({link]a). Each of these globin molecules is bound to a red 
pigment molecule called heme, which contains an ion of iron (Fe**) 
([link]b). 

Hemoglobin 


B chain 1 B chain 2 


a chain 1 


(a) A molecule of hemoglobin contains four globin 
proteins, each of which is bound to one molecule of the 
iron-containing pigment heme. (b) A single erythrocyte 
can contain 300 million hemoglobin molecules, and thus 

more than 1 billion oxygen molecules. 


Each iron ion in the heme can bind to one oxygen molecule; therefore, each 
hemoglobin molecule can transport four oxygen molecules. An individual 
erythrocyte may contain about 300 million hemoglobin molecules, and 
therefore can bind to and transport up to 1.2 billion oxygen molecules (see 
[link]b). 


In the lungs, hemoglobin picks up oxygen, which binds to the iron ions, 
forming oxyhemoglobin. The bright red, oxygenated hemoglobin travels to 
the body tissues, where it releases some of the oxygen molecules, becoming 
darker red deoxyhemoglobin, sometimes referred to as reduced 
hemoglobin. Oxygen release depends on the need for oxygen in the 
surrounding tissues, so hemoglobin rarely if ever leaves all of its oxygen 
behind. In the capillaries, carbon dioxide enters the bloodstream. About 76 
percent dissolves in the plasma, some of it remaining as dissolved CO>, and 
the remainder forming bicarbonate ion. About 23—24 percent of it binds to 
the amino acids in hemoglobin, forming a molecule known as 
carbaminohemoglobin. From the capillaries, the hemoglobin carries 


carbon dioxide back to the lungs, where it releases it for exchange of 
oxygen. 


Changes in the levels of RBCs can have significant effects on the body’s 
ability to effectively deliver oxygen to the tissues. Ineffective hematopoiesis 
results in insufficient numbers of RBCs and results in one of several forms 
of anemia. An overproduction of RBCs produces a condition called 
polycythemia. The primary drawback with polycythemia is not a failure to 
directly deliver enough oxygen to the tissues, but rather the increased 
viscosity of the blood, which makes it more difficult for the heart to 
circulate the blood. 


In patients with insufficient hemoglobin, the tissues may not receive 
sufficient oxygen, resulting in another form of anemia. In determining 
oxygenation of tissues, the value of greatest interest in healthcare is the 
percent saturation; that is, the percentage of hemoglobin sites occupied by 
oxygen in a patient’s blood. Clinically this value is commonly referred to 
simply as “percent sat.” 


Percent saturation is normally monitored using a device known as a pulse 
oximeter, which is applied to a thin part of the body, typically the tip of the 
patient’s finger. The device works by sending two different wavelengths of 
light (one red, the other infrared) through the finger and measuring the light 
with a photodetector as it exits. Hemoglobin absorbs light differentially 
depending upon its saturation with oxygen. The machine calibrates the 
amount of light received by the photodetector against the amount absorbed 
by the partially oxygenated hemoglobin and presents the data as percent 
saturation. Normal pulse oximeter readings range from 95-100 percent. 
Lower percentages reflect hypoxemia, or low blood oxygen. The term 
hypoxia is more generic and simply refers to low oxygen levels. Oxygen 
levels are also directly monitored from free oxygen in the plasma typically 
following an arterial stick. When this method is applied, the amount of 
oxygen present is expressed in terms of partial pressure of oxygen or simply 
pO, and is typically recorded in units of millimeters of mercury, mm Hg. 


The kidneys filter about 180 liters (~380 pints) of blood in an average adult 
each day, or about 20 percent of the total resting volume, and thus serve as 
ideal sites for receptors that determine oxygen saturation. In response to 


hypoxemia, less oxygen will exit the vessels supplying the kidney, resulting 
in hypoxia (low oxygen concentration) in the tissue fluid of the kidney 
where oxygen concentration is actually monitored. Interstitial fibroblasts 
within the kidney secrete EPO, thereby increasing erythrocyte production 
and restoring oxygen levels. In a classic negative-feedback loop, as oxygen 
saturation rises, EPO secretion falls, and vice versa, thereby maintaining 
homeostasis. Populations dwelling at high elevations, with inherently lower 
levels of oxygen in the atmosphere, naturally maintain a hematocrit higher 
than people living at sea level. Consequently, people traveling to high 
elevations may experience symptoms of hypoxemia, such as fatigue, 
headache, and shortness of breath, for a few days after their arrival. In 
response to the hypoxemia, the kidneys secrete EPO to step up the 
production of erythrocytes until homeostasis is achieved once again. To 
avoid the symptoms of hypoxemia, or altitude sickness, mountain climbers 
typically rest for several days to a week or more at a series of camps 
situated at increasing elevations to allow EPO levels and, consequently, 
erythrocyte counts to rise. When climbing the tallest peaks, such as Mt. 
Everest and K2 in the Himalayas, many mountain climbers rely upon 
bottled oxygen as they near the summit. 


Lifecycle of Erythrocytes 


Production of erythrocytes in the marrow occurs at the staggering rate of 
more than 2 million cells per second. For this production to occur, a number 
of raw materials must be present in adequate amounts. These include the 
same nutrients that are essential to the production and maintenance of any 
cell, such as glucose, lipids, and amino acids. However, erythrocyte 
production also requires several trace elements: 


e Iron. We have said that each heme group in a hemoglobin molecule 
contains an ion of the trace mineral iron. On average, less than 20 
percent of the iron we consume is absorbed. Heme iron, from animal 
foods such as meat, poultry, and fish, is absorbed more efficiently than 
non-heme iron from plant foods. Upon absorption, iron becomes part 
of the body’s total iron pool. The bone marrow, liver, and spleen can 
store iron in the protein compounds ferritin and hemosiderin. 
Ferroportin transports the iron across the intestinal cell plasma 


membranes and from its storage sites into tissue fluid where it enters 
the blood. When EPO stimulates the production of erythrocytes, iron is 
released from storage, bound to transferrin, and carried to the red 
marrow where it attaches to erythrocyte precursors. 

e Copper. A trace mineral, copper is a component of two plasma 
proteins, hephaestin and ceruloplasmin. Without these, hemoglobin 
could not be adequately produced. Located in intestinal villi, 
hephaestin enables iron to be absorbed by intestinal cells. 
Ceruloplasmin transports copper. Both enable the oxidation of iron 
from Fe** to Fe**, a form in which it can be bound to its transport 
protein, transferrin, for transport to body cells. In a state of copper 
deficiency, the transport of iron for heme synthesis decreases, and iron 
can accumulate in tissues, where it can eventually lead to organ 
damage. 

e Zinc. The trace mineral zinc functions as a co-enzyme that facilitates 
the synthesis of the heme portion of hemoglobin. 

e B vitamins. The B vitamins folate and vitamin B,5 function as co- 
enzymes that facilitate DNA synthesis. Thus, both are critical for the 
synthesis of new cells, including erythrocytes. 


Erythrocytes live up to 120 days in the circulation, after which the worn-out 
cells are removed by a type of myeloid phagocytic cell called a 
macrophage, located primarily within the bone marrow, liver, and spleen. 
The components of the degraded erythrocytes’ hemoglobin are further 
processed as follows: 


¢ Globin, the protein portion of hemoglobin, is broken down into amino 
acids, which can be sent back to the bone marrow to be used in the 
production of new erythrocytes. Hemoglobin that is not phagocytized 
is broken down in the circulation, releasing alpha and beta chains that 
are removed from circulation by the kidneys. 

e The iron contained in the heme portion of hemoglobin may be stored 
in the liver or spleen, primarily in the form of ferritin or hemosiderin, 
or carried through the bloodstream by transferrin to the red bone 
marrow for recycling into new erythrocytes. 

e The non-iron portion of heme is degraded into the waste product 
biliverdin, a green pigment, and then into another waste product, 


bilirubin, a yellow pigment. Bilirubin binds to albumin and travels in 
the blood to the liver, which uses it in the manufacture of bile, a 
compound released into the intestines to help emulsify dietary fats. In 
the large intestine, bacteria breaks the bilirubin apart from the bile and 
converts it to urobilinogen and then into stercobilin. It is then 
eliminated from the body in the feces. Broad-spectrum antibiotics 
typically eliminate these bacteria as well and may alter the color of 
feces. The kidneys also remove any circulating bilirubin and other 
related metabolic byproducts such as urobilins and secrete them into 
the urine. 


The breakdown pigments formed from the destruction of hemoglobin can 
be seen in a variety of situations. At the site of an injury, biliverdin from 
damaged RBCs produces some of the dramatic colors associated with 
bruising. With a failing liver, bilirubin cannot be removed effectively from 
circulation and causes the body to assume a yellowish tinge associated with 
jaundice. Stercobilins within the feces produce the typical brown color 
associated with this waste. And the yellow of urine is associated with the 
urobilins. 


The erythrocyte lifecycle is summarized in [link]. 
Erythrocyte Lifecycle 


Hemopoiesis of erythrocytes begins 


©) Unused heme groups can be recycled and used in in the hemopoietic bone marrow. 


hemopoiesis, or can be converted into bilirubin 
and used to make bile in the liver. Iron ions 
can also be transferred to the protein ferritin 
for storage in the liver. 


Locations of hemopoietic 
bone marrow 


@ Stem cell 
@ Erythroblast 


Bilirubin Ferritin 


Iron ions 
Biliverdin bound 
to transferrin 


©) The heme portion 
is broken down into 
biliverdin for 
transport in the 
blood. The iron ions 
bind to the blood 
protein transferrin 
for transport. 


Globin 
Heme oR ihS amino acids 
groups and cell @) Reticulocytes are released into the 


bloodstream, where they mature into 
erythrocytes, which circulate for an 
average of 120 days. 


\/ components 


Hemoglobin 
protein 
structure 


@) Old and damaged 
erythrocytes are 
phagocytized by 


is broken macrophages in 
down into the bone marrow, 
amino acids liver, and spleen. 


Lysosome 


@ The globin (protein) portion of hemoglobin 
is metabolized into amino acids, which are 
reused for protein synthesis. 


Erythrocytes are produced in the bone marrow and sent into the 
circulation. At the end of their lifecycle, they are destroyed by 


macrophages, and their components are recycled. 


Disorders of Erythrocytes 


The size, shape, and number of erythrocytes, and the number of hemoglobin 
molecules can have a major impact on a person’s health. When the number 
of RBCs or hemoglobin is deficient, the general condition is called anemia. 
There are more than 400 types of anemia and more than 3.5 million 
Americans suffer from this condition. Anemia can be broken down into 
three major groups: those caused by blood loss, those caused by faulty or 
decreased RBC production, and those caused by excessive destruction of 
RBCs. Clinicians often use two groupings in diagnosis: The kinetic 
approach focuses on evaluating the production, destruction, and removal of 
RBCs, whereas the morphological approach examines the RBCs 
themselves, paying particular emphasis to their size. A common test is the 
mean corpuscle volume (MCV), which measures size. Normal-sized cells 
are referred to as normocytic, smaller-than-normal cells are referred to as 
microcytic, and larger-than-normal cells are referred to as macrocytic. 
Reticulocyte counts are also important and may reveal inadequate 
production of RBCs. The effects of the various anemias are widespread, 
because reduced numbers of RBCs or hemoglobin will result in lower levels 
of oxygen being delivered to body tissues. Since oxygen is required for 
tissue functioning, anemia produces fatigue, lethargy, and an increased risk 
for infection. An oxygen deficit in the brain impairs the ability to think 
clearly, and may prompt headaches and irritability. Lack of oxygen leaves 
the patient short of breath, even as the heart and lungs work harder in 
response to the deficit. 


Blood loss anemias are fairly straightforward. In addition to bleeding from 
wounds or other lesions, these forms of anemia may be due to ulcers, 
hemorrhoids, inflammation of the stomach (gastritis), and some cancers of 
the gastrointestinal tract. The excessive use of aspirin or other nonsteroidal 
anti-inflammatory drugs such as ibuprofen can trigger ulceration and 
gastritis. Excessive menstruation and loss of blood during childbirth are 
also potential causes. 


Anemias caused by faulty or decreased RBC production include sickle cell 
anemia, iron deficiency anemia, vitamin deficiency anemia, and diseases of 
the bone marrow and stem cells. 


e A characteristic change in the shape of erythrocytes is seen in sickle 
cell disease (also referred to as sickle cell anemia). A genetic disorder, 
it is caused by production of an abnormal type of hemoglobin, called 
hemoglobin S, which delivers less oxygen to tissues and causes 
erythrocytes to assume a sickle (or crescent) shape, especially at low 
oxygen concentrations ({link]). These abnormally shaped cells can 
then become lodged in narrow capillaries because they are unable to 
fold in on themselves to squeeze through, blocking blood flow to 
tissues and causing a variety of serious problems from painful joints to 
delayed growth and even blindness and cerebrovascular accidents 
(strokes). Sickle cell anemia is a genetic condition particularly found 
in individuals of African descent. 


Sickle Cells 


Sickle cell anemia is caused by 
a mutation in one of the 
hemoglobin genes. Erythrocytes 
produce an abnormal type of 


hemoglobin, which causes the 
cell to take on a sickle or 
crescent shape. (credit: Janice 
Haney Carr) 


e Iron deficiency anemia is the most common type and results when the 
amount of available iron is insufficient to allow production of 
sufficient heme. This condition can occur in individuals with a 
deficiency of iron in the diet and is especially common in teens and 
children as well as in vegans and vegetarians. Additionally, iron 
deficiency anemia may be caused by either an inability to absorb and 
transport iron or slow, chronic bleeding. 

¢ Vitamin-deficient anemias generally involve insufficient vitamin B12 
and folate. 


o Megaloblastic anemia involves a deficiency of vitamin B12 
and/or folate, and often involves diets deficient in these essential 
nutrients. Lack of meat or a viable alternate source, and 
overcooking or eating insufficient amounts of vegetables may 
lead to a lack of folate. 

o Pernicious anemia is caused by poor absorption of vitamin B12 
and is often seen in patients with Crohn’s disease (a severe 
intestinal disorder often treated by surgery), surgical removal of 
the intestines or stomach (common in some weight loss 
surgeries), intestinal parasites, and AIDS. 

o Pregnancies, some medications, excessive alcohol consumption, 
and some diseases such as celiac disease are also associated with 
vitamin deficiencies. It is essential to provide sufficient folic acid 
during the early stages of pregnancy to reduce the risk of 
neurological defects, including spina bifida, a failure of the neural 
tube to close. 


e Assorted disease processes can also interfere with the production and 
formation of RBCs and hemoglobin. If myeloid stem cells are 


defective or replaced by cancer cells, there will be insufficient 
quantities of RBCs produced. 


o Aplastic anemia is the condition in which there are deficient 
numbers of RBC stem cells. Aplastic anemia is often inherited, or 
it may be triggered by radiation, medication, chemotherapy, or 
infection. 

o Thalassemia is an inherited condition typically occurring in 
individuals from the Middle East, the Mediterranean, African, and 
Southeast Asia, in which maturation of the RBCs does not 
proceed normally. The most severe form is called Cooley’s 
anemia. 

o Lead exposure from industrial sources or even dust from paint 
chips of iron-containing paints or pottery that has not been 
properly glazed may also lead to destruction of the red marrow. 


e Various disease processes also can lead to anemias. These include 
chronic kidney diseases often associated with a decreased production 
of EPO, hypothyroidism, some forms of cancer, lupus, and rheumatoid 
arthritis. 


In contrast to anemia, an elevated RBC count is called polycythemia and is 
detected in a patient’s elevated hematocrit. It can occur transiently in a 
person who is dehydrated; when water intake is inadequate or water losses 
are excessive, the plasma volume falls. As a result, the hematocrit rises. For 
reasons mentioned earlier, a mild form of polycythemia is chronic but 
normal in people living at high altitudes. Some elite athletes train at high 
elevations specifically to induce this phenomenon. Finally, a type of bone 
marrow disease called polycythemia vera (from the Greek vera = “true”) 
causes an excessive production of immature erythrocytes. Polycythemia 
vera can dangerously elevate the viscosity of blood, raising blood pressure 
and making it more difficult for the heart to pump blood throughout the 
body. It is a relatively rare disease that occurs more often in men than 
women, and is more likely to be present in elderly patients those over 60 
years of age. 


Chapter Review 


The most abundant formed elements in blood, erythrocytes are red, 
biconcave disks packed with an oxygen-carrying compound called 
hemoglobin. The hemoglobin molecule contains four globin proteins bound 
to a pigment molecule called heme, which contains an ion of iron. In the 
bloodstream, iron picks up oxygen in the lungs and drops it off in the 
tissues; the amino acids in hemoglobin then transport carbon dioxide from 
the tissues back to the lungs. Erythrocytes live only 120 days on average, 
and thus must be continually replaced. Worn-out erythrocytes are 
phagocytized by macrophages and their hemoglobin is broken down. The 
breakdown products are recycled or removed as wastes: Globin is broken 
down into amino acids for synthesis of new proteins; iron is stored in the 
liver or spleen or used by the bone marrow for production of new 
erythrocytes; and the remnants of heme are converted into bilirubin, or 
other waste products that are taken up by the liver and excreted in the bile 
or removed by the kidneys. Anemia is a deficiency of RBCs or hemoglobin, 
whereas polycythemia is an excess of RBCs. 


Review Questions 


Exercise: 


Problem: 


Which of the following statements about mature, circulating 
erythrocytes is true? 


a. They have no nucleus. 

b. They are packed with mitochondria. 

c. They survive for an average of 4 days. 
d. All of the above 


Solution: 


A 


Exercise: 


Problem:A molecule of hemoglobin 


a. is shaped like a biconcave disk packed almost entirely with iron 

b. contains four glycoprotein units studded with oxygen 

c. consists of four globin proteins, each bound to a molecule of 
heme 

d. can carry up to 120 molecules of oxygen 


Solution: 


C 
Exercise: 


Problem: 


The production of healthy erythrocytes depends upon the availability 
of 


a. Copper 
b. zinc 

c. vitamin By 

d. copper, zinc, and vitamin By 


Solution: 


D 
Exercise: 


Problem: 


Aging and damaged erythrocytes are removed from the circulation by 


a. myeoblasts 
b. monocytes 


c. macrophages 
d. mast cells 


Solution: 


C 
Exercise: 


Problem: 


A patient has been suffering for 2 months with a chronic, watery 
diarrhea. A blood test is likely to reveal 


a. a hematocrit below 30 percent 
b. hypoxemia 

c. anemia 

d. polycythemia 


Solution: 


D 


Critical Thinking Questions 


Exercise: 
Problem: 
A young woman has been experiencing unusually heavy menstrual 
bleeding for several years. She follows a strict vegan diet (no animal 
foods). She is at risk for what disorder, and why? 


Solution: 


She is at risk for anemia, because her unusually heavy menstrual 
bleeding results in excessive loss of erythrocytes each month. At the 


same time, her vegan diet means that she does not have dietary sources 
of heme iron. The non-heme iron she consumes in plant foods is not as 
well absorbed as heme iron. 


Exercise: 


Problem: 


A patient has thalassemia, a genetic disorder characterized by 
abnormal synthesis of globin proteins and excessive destruction of 
erythrocytes. This patient is jaundiced and is found to have an 
excessive level of bilirubin in his blood. Explain the connection. 


Solution: 


Bilirubin is a breakdown product of the non-iron component of heme, 
which is cleaved from globin when erythrocytes are degraded. 
Excessive erythrocyte destruction would deposit excessive bilirubin in 
the blood. Bilirubin is a yellowish pigment, and high blood levels can 
manifest as yellowed skin. 


Glossary 


anemia 
deficiency of red blood cells or hemoglobin 


bilirubin 
yellowish bile pigment produced when iron is removed from heme and 
is further broken down into waste products 


biliverdin 
green bile pigment produced when the non-iron portion of heme is 
degraded into a waste product; converted to bilirubin in the liver 


carbaminohemoglobin 
compound of carbon dioxide and hemoglobin, and one of the ways in 
which carbon dioxide is carried in the blood 


deoxyhemoglobin 
molecule of hemoglobin without an oxygen molecule bound to it 


erythrocyte 
(also, red blood cell) mature myeloid blood cell that is composed 
mostly of hemoglobin and functions primarily in the transportation of 
oxygen and carbon dioxide 


ferritin 
protein-containing storage form of iron found in the bone marrow, 
liver, and spleen 


globin 
heme-containing globular protein that is a constituent of hemoglobin 


heme 
red, iron-containing pigment to which oxygen binds in hemoglobin 


hemoglobin 
oxygen-carrying compound in erythrocytes 


hemosiderin 
protein-containing storage form of iron found in the bone marrow, 
liver, and spleen 


hypoxemia 
below-normal level of oxygen saturation of blood (typically <95 
percent) 


macrophage 
phagocytic cell of the myeloid lineage; a matured monocyte 


oxyhemoglobin 
molecule of hemoglobin to which oxygen is bound 


polycythemia 
elevated level of hemoglobin, whether adaptive or pathological 


reticulocyte 


immature erythrocyte that may still contain fragments of organelles 


sickle cell disease 
(also, sickle cell anemia) inherited blood disorder in which 
hemoglobin molecules are malformed, leading to the breakdown of 
RBCs that take on a characteristic sickle shape 


thalassemia 
inherited blood disorder in which maturation of RBCs does not 
proceed normally, leading to abnormal formation of hemoglobin and 
the destruction of RBCs 


transferrin 
plasma protein that binds reversibly to iron and distributes it 
throughout the body 


Leukocytes and Platelets 
By the end of this section, you will be able to: 


e Describe the general characteristics of leukocytes 

¢ Classify leukocytes according to their lineage, their main structural 
features, and their primary functions 

e Discuss the most common malignancies involving leukocytes 

e Identify the lineage, basic structure, and function of platelets 


The leukocyte, commonly known as a white blood cell (or WBC), is a 
major component of the body’s defenses against disease. Leukocytes 
protect the body against invading microorganisms and body cells with 
mutated DNA, and they clean up debris. Platelets are essential for the repair 
of blood vessels when damage to them has occurred; they also provide 
growth factors for healing and repair. See [link] for a summary of 
leukocytes and platelets. 


Characteristics of Leukocytes 


Although leukocytes and erythrocytes both originate from hematopoietic 
stem cells in the bone marrow, they are very different from each other in 
many significant ways. For instance, leukocytes are far less numerous than 
erythrocytes: Typically there are only 5000 to 10,000 per pL. They are also 
larger than erythrocytes and are the only formed elements that are complete 
cells, possessing a nucleus and organelles. And although there is just one 
type of erythrocyte, there are many types of leukocytes. Most of these types 
have a much shorter lifespan than that of erythrocytes, some as short as a 
few hours or even a few minutes in the case of acute infection. 


One of the most distinctive characteristics of leukocytes is their movement. 
Whereas erythrocytes spend their days circulating within the blood vessels, 
leukocytes routinely leave the bloodstream to perform their defensive 
functions in the body’s tissues. For leukocytes, the vascular network is 
simply a highway they travel and soon exit to reach their true destination. 
When they arrive, they are often given distinct names, such as macrophage 
or microglia, depending on their function. As shown in [link], they leave the 
capillaries—the smallest blood vessels—or other small vessels through a 


process known as emigration (from the Latin for “removal”) or diapedesis 
(dia- = “through”; -pedan = “to leap”) in which they squeeze through 
adjacent cells in a blood vessel wall. 


Once they have exited the capillaries, some leukocytes will take up fixed 
positions in lymphatic tissue, bone marrow, the spleen, the thymus, or other 
organs. Others will move about through the tissue spaces very much like 
amoebas, continuously extending their plasma membranes, sometimes 
wandering freely, and sometimes moving toward the direction in which they 
are drawn by chemical signals. This attracting of leukocytes occurs because 
of positive chemotaxis (literally “movement in response to chemicals”), a 
phenomenon in which injured or infected cells and nearby leukocytes emit 
the equivalent of a chemical “911” call, attracting more leukocytes to the 
site. In clinical medicine, the differential counts of the types and 
percentages of leukocytes present are often key indicators in making a 
diagnosis and selecting a treatment. 

Emigration 


GQ) Leukocytes in the blood 
respond to chemical Eosinophil 
attractants released by 
pathogens and 


Injured/infected cells 
secrete chemical signals 


chemical signals from into the blood. 
nearby injured cells. 
Monocyte 
Neutrophil : Pathogens 
Leukocytes emigrate 
Q@) The leukocytes squeeze to site of injury and 
between the cells of infection. 
the capillary wall as they 
follow the chemical 
signals to where they 
are most concentrated 
(positive chemotaxis). 
Eosinophil releases 
cytotoxic chemicals 
from granules into 
tissue. 
@) Within the damaged tissue, 


monocytes differentiate into 
macrophages that phagocytize the 
pathogens. The eosinophils and 
neutrophils release chemicals that 
break apart pathogens. They are 
also capable of phagocytosis. 
Macrophage engulfs 
pathogen. 


Leukocytes exit the blood vessel and then move through the 
connective tissue of the dermis toward the site of a wound. Some 
leukocytes, such as the eosinophil and neutrophil, are characterized 


as granular leukocytes. They release chemicals from their granules 

that destroy pathogens; they are also capable of phagocytosis. The 

monocyte, an agranular leukocyte, differentiates into a macrophage 
that then phagocytizes the pathogens. 


Classification of Leukocytes 


When scientists first began to observe stained blood slides, it quickly 
became evident that leukocytes could be divided into two groups, according 
to whether their cytoplasm contained highly visible granules: 


¢ Granular leukocytes contain abundant granules within the cytoplasm. 
They include neutrophils, eosinophils, and basophils (you can view 
their lineage from myeloid stem cells in [link]). 

e While granules are not totally lacking in agranular leukocytes, they 
are far fewer and less obvious. Agranular leukocytes include 
monocytes, which mature into macrophages that are phagocytic, and 
lymphocytes, which arise from the lymphoid stem cell line. 


Granular Leukocytes 


We will consider the granular leukocytes in order from most common to 
least common. All of these are produced in the red bone marrow and have a 
short lifespan of hours to days. They typically have a lobed nucleus and are 
classified according to which type of stain best highlights their granules 
({link]). 

Granular Leukocytes 


és 


Neutrophil Eosinophil Basophil 


A neutrophil has small granules that stain 
light lilac and a nucleus with two to five 
lobes. An eosinophil’s granules are 
slightly larger and stain reddish-orange, 
and its nucleus has two to three lobes. A 
basophil has large granules that stain dark 
blue to purple and a two-lobed nucleus. 


The most common of all the leukocytes, neutrophils will normally 
comprise 50—70 percent of total leukocyte count. They are 10-12 pm in 
diameter, significantly larger than erythrocytes. They are called neutrophils 
because their granules show up most clearly with stains that are chemically 
neutral (neither acidic nor basic). The granules are numerous but quite fine 
and normally appear light lilac. The nucleus has a distinct lobed appearance 
and may have two to five lobes, the number increasing with the age of the 
cell. Older neutrophils have increasing numbers of lobes and are often 
referred to as polymorphonuclear (a nucleus with many forms), or simply 
“polys.” Younger and immature neutrophils begin to develop lobes and are 
known as “bands.” 


Neutrophils are rapid responders to the site of infection and are efficient 
phagocytes with a preference for bacteria. Their granules include lysozyme, 
an enzyme capable of lysing, or breaking down, bacterial cell walls; 
oxidants such as hydrogen peroxide; and defensins, proteins that bind to 
and puncture bacterial and fungal plasma membranes, so that the cell 
contents leak out. Abnormally high counts of neutrophils indicate infection 
and/or inflammation, particularly triggered by bacteria, but are also found 
in burn patients and others experiencing unusual stress. A burn injury 
increases the proliferation of neutrophils in order to fight off infection that 
can result from the destruction of the barrier of the skin. Low counts may be 
caused by drug toxicity and other disorders, and may increase an 
individual’s susceptibility to infection. 


Eosinophils typically represent 2—4 percent of total leukocyte count. They 
are also 10—12 pm in diameter. The granules of eosinophils stain best with 


an acidic stain known as eosin. The nucleus of the eosinophil will typically 
have two to three lobes and, if stained properly, the granules will have a 
distinct red to orange color. 


The granules of eosinophils include antihistamine molecules, which 
counteract the activities of histamines, inflammatory chemicals produced by 
basophils and mast cells. Some eosinophil granules contain molecules toxic 
to parasitic worms, which can enter the body through the integument, or 
when an individual consumes raw or undercooked fish or meat. Eosinophils 
are also capable of phagocytosis and are particularly effective when 
antibodies bind to the target and form an antigen-antibody complex. High 
counts of eosinophils are typical of patients experiencing allergies, parasitic 
wor! infestations, and some autoimmune diseases. Low counts may be due 
to drug toxicity and stress. 


Basophils are the least common leukocytes, typically comprising less than 
one percent of the total leukocyte count. They are slightly smaller than 
neutrophils and eosinophils at 8-10 yum in diameter. The granules of 
basophils stain best with basic (alkaline) stains. Basophils contain large 
granules that pick up a dark blue stain and are so common they may make it 
difficult to see the two-lobed nucleus. 


In general, basophils intensify the inflammatory response. They share this 
trait with mast cells. In the past, mast cells were considered to be basophils 
that left the circulation. However, this appears not to be the case, as the two 
cell types develop from different lineages. 


The granules of basophils release histamines, which contribute to 
inflammation, and heparin, which opposes blood clotting. High counts of 
basophils are associated with allergies, parasitic infections, and 
hypothyroidism. Low counts are associated with pregnancy, stress, and 
hyperthyroidism. 


Agranular Leukocytes 


Agranular leukocytes contain smaller, less-visible granules in their 

cytoplasm than do granular leukocytes. The nucleus is simple in shape, 
sometimes with an indentation but without distinct lobes. There are two 
major types of agranulocytes: lymphocytes and monocytes (see [link]). 


Lymphocytes are the only formed element of blood that arises from 
lymphoid stem cells. Although they form initially in the bone marrow, 
much of their subsequent development and reproduction occurs in the 
lymphatic tissues. Lymphocytes are the second most common type of 
leukocyte, accounting for about 20—30 percent of all leukocytes, and are 
essential for the immune response. The size range of lymphocytes is quite 
extensive, with some authorities recognizing two size classes and others 
three. Typically, the large cells are 10-14 ym and have a smaller nucleus-to- 
cytoplasm ratio and more granules. The smaller cells are typically 6-9 pm 
with a larger volume of nucleus to cytoplasm, creating a “halo” effect. A 
few cells may fall outside these ranges, at 14-17 pm. This finding has led to 
the three size range classification. 


The three major groups of lymphocytes include natural killer cells, B cells, 
and T cells. Natural killer (NK) cells are capable of recognizing cells that 
do not express “self” proteins on their plasma membrane or that contain 
foreign or abnormal markers. These “nonself” cells include cancer cells, 
cells infected with a virus, and other cells with atypical surface proteins. 
Thus, they provide generalized, nonspecific immunity. The larger 
lymphocytes are typically NK cells. 


B cells and T cells, also called B lymphocytes and T lymphocytes, play 
prominent roles in defending the body against specific pathogens (disease- 
causing microorganisms) and are involved in specific immunity. One form 
of B cells (plasma cells) produces the antibodies or immunoglobulins that 
bind to specific foreign or abnormal components of plasma membranes. 
This is also referred to as humoral (body fluid) immunity. T cells provide 
cellular-level immunity by physically attacking foreign or diseased cells. A 
memory cell is a variety of both B and T cells that forms after exposure to 
a pathogen and mounts rapid responses upon subsequent exposures. Unlike 
other leukocytes, memory cells live for many years. B cells undergo a 
maturation process in the bone marrow, whereas T cells undergo maturation 


in the thymus. This site of the maturation process gives rise to the name B 
and T cells. The functions of lymphocytes are complex and will be covered 
in detail in the chapter covering the lymphatic system and immunity. 
Smaller lymphocytes are either B or T cells, although they cannot be 
differentiated in a normal blood smear. 


Abnormally high lymphocyte counts are characteristic of viral infections as 
well as some types of cancer. Abnormally low lymphocyte counts are 
characteristic of prolonged (chronic) illness or immunosuppression, 
including that caused by HIV infection and drug therapies that often involve 
steroids. 


Monocytes originate from myeloid stem cells. They normally represent 2—8 
percent of the total leukocyte count. They are typically easily recognized by 
their large size of 12—20 ym and indented or horseshoe-shaped nuclei. 
Macrophages are monocytes that have left the circulation and phagocytize 
debris, foreign pathogens, worn-out erythrocytes, and many other dead, 
worn out, or damaged cells. Macrophages also release antimicrobial 
defensins and chemotactic chemicals that attract other leukocytes to the site 
of an infection. Some macrophages occupy fixed locations, whereas others 
wander through the tissue fluid. 


Abnormally high counts of monocytes are associated with viral or fungal 
infections, tuberculosis, and some forms of leukemia and other chronic 
diseases. Abnormally low counts are typically caused by suppression of the 
bone marrow. 


Lifecycle of Leukocytes 


Most leukocytes have a relatively short lifespan, typically measured in 
hours or days. Production of all leukocytes begins in the bone marrow 
under the influence of CSFs and interleukins. Secondary production and 
maturation of lymphocytes occurs in specific regions of lymphatic tissue 
known as germinal centers. Lymphocytes are fully capable of mitosis and 
may produce clones of cells with identical properties. This capacity enables 
an individual to maintain immunity throughout life to many threats that 
have been encountered in the past. 


Disorders of Leukocytes 


Leukopenia is a condition in which too few leukocytes are produced. If this 
condition is pronounced, the individual may be unable to ward off disease. 
Excessive leukocyte proliferation is known as leukocytosis. Although 
leukocyte counts are high, the cells themselves are often nonfunctional, 
leaving the individual at increased risk for disease. 


Leukemia is a cancer involving an abundance of leukocytes. It may involve 
only one specific type of leukocyte from either the myeloid line (myelocytic 
leukemia) or the lymphoid line (lymphocytic leukemia). In chronic 
leukemia, mature leukocytes accumulate and fail to die. In acute leukemia, 
there is an overproduction of young, immature leukocytes. In both 
conditions the cells do not function properly. 


Lymphoma is a form of cancer in which masses of malignant T and/or B 
lymphocytes collect in lymph nodes, the spleen, the liver, and other tissues. 
As in leukemia, the malignant leukocytes do not function properly, and the 
patient is vulnerable to infection. Some forms of lymphoma tend to 
progress slowly and respond well to treatment. Others tend to progress 
quickly and require aggressive treatment, without which they are rapidly 
fatal. 


Platelets 


You may occasionally see platelets referred to as thrombocytes, but 
because this name suggests they are a type of cell, it is not accurate. A 
platelet is not a cell but rather a fragment of the cytoplasm of a cell called a 
megakaryocyte that is surrounded by a plasma membrane. Megakaryocytes 
are descended from myeloid stem cells (see [link]) and are large, typically 
50-100 pm in diameter, and contain an enlarged, lobed nucleus. As noted 
earlier, thrombopoietin, a glycoprotein secreted by the kidneys and liver, 
stimulates the proliferation of megakaryoblasts, which mature into 
megakaryocytes. These remain within bone marrow tissue ([link]) and 
ultimately form platelet-precursor extensions that extend through the walls 
of bone matrow capillaries to release into the circulation thousands of 
cytoplasmic fragments, each enclosed by a bit of plasma membrane. These 


enclosed fragments are platelets. Each megakarocyte releases 2000—3000 
platelets during its lifespan. Following platelet release, megakaryocyte 
remnants, which are little more than a cell nucleus, are consumed by 
macrophages. 


Platelets are relatively small, 2-4 ym in diameter, but numerous, with 
typically 150,000—160,000 per uL of blood. After entering the circulation, 
approximately one-third migrate to the spleen for storage for later release in 
response to any rupture in a blood vessel. They then become activated to 
perform their primary function, which is to limit blood loss. Platelets 
remain only about 10 days, then are phagocytized by macrophages. 


Platelets are critical to hemostasis, the stoppage of blood flow following 
damage to a vessel. They also secrete a variety of growth factors essential 
for growth and repair of tissue, particularly connective tissue. Infusions of 
concentrated platelets are now being used in some therapies to stimulate 
healing. 


Disorders of Platelets 


Thrombocytosis is a condition in which there are too many platelets. This 
may trigger formation of unwanted blood clots (thrombosis), a potentially 
fatal disorder. If there is an insufficient number of platelets, called 
thrombocytopenia, blood may not clot properly, and excessive bleeding 
may result. 

Platelets 


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Leukocytes 


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Basophil Eosinophil Neutrophil Monocyte Lymphocyte 


(Micrographs provided by the Regents of 


University of Michigan Medical School © 
2012) 


View University of Michigan Webscopes at 
http://virtualslides.med.umich.edu/Histology/Cardiovascular%20System/0 
81-2 HISTO 40X.svs/view.apml? 
cwidth=860&cheight=733&chost=virtualslides.med.umich.edu&dlistview= 
1&title=&csis=1 and explore the blood slides in greater detail. The 
Webscope feature allows you to move the slides as you would with a 
mechanical stage. You can increase and decrease the magnification. There 
is a chance to review each of the leukocytes individually after you have 
attempted to identify them from the first two blood smears. In addition, 
there are a few multiple choice questions. 

Are you able to recognize and identify the various formed elements? You 
will need to do this is a systematic manner, scanning along the image. The 
standard method is to use a grid, but this is not possible with this resource. 
Try constructing a simple table with each leukocyte type and then making a 
mark for each cell type you identify. Attempt to classify at least 50 and 
perhaps as many as 100 different cells. Based on the percentage of cells 
that you count, do the numbers represent a normal blood smear or does 
something appear to be abnormal? 


Chapter Review 


Leukocytes function in body defenses. They squeeze out of the walls of 
blood vessels through emigration or diapedesis, then may move through 
tissue fluid or become attached to various organs where they fight against 
pathogenic organisms, diseased cells, or other threats to health. Granular 
leukocytes, which include neutrophils, eosinophils, and basophils, originate 
with myeloid stem cells, as do the agranular monocytes. The other 
agranular leukocytes, NK cells, B cells, and T cells, arise from the 
lymphoid stem cell line. The most abundant leukocytes are the neutrophils, 
which are first responders to infections, especially with bacteria. About 20— 
30 percent of all leukocytes are lymphocytes, which are critical to the 


body’s defense against specific threats. Leukemia and lymphoma are 
malignancies involving leukocytes. Platelets are fragments of cells known 
as megakaryocytes that dwell within the bone marrow. While many 
platelets are stored in the spleen, others enter the circulation and are 
essential for hemostasis; they also produce several growth factors important 
for repair and healing. 


Interactive Link Questions 


Exercise: 


Problem: 


[link] Are you able to recognize and identify the various formed 
elements? You will need to do this is a systematic manner, scanning 
along the image. The standard method is to use a grid, but this is not 
possible with this resource. Try constructing a simple table with each 
leukocyte type and then making a mark for each cell type you identify. 
Attempt to classify at least 50 and perhaps as many as 100 different 
cells. Based on the percentage of cells that you count, do the numbers 
represent a normal blood smear or does something appear to be 
abnormal? 


Solution: 


[link] This should appear to be a normal blood smear. 


Review Questions 


Exercise: 


Problem: 


The process by which leukocytes squeeze through adjacent cells in a 
blood vessel wall is called 


a. leukocytosis 
b. positive chemotaxis 


c. emigration 
d. cytoplasmic extending 


Solution: 


CG 


Exercise: 


Problem: Which of the following describes a neutrophil? 


a. abundant, agranular, especially effective against cancer cells 

b. abundant, granular, especially effective against bacteria 

c. rare, agranular, releases antimicrobial defensins 

d. rare, granular, contains multiple granules packed with histamine 


Solution: 


B 


Exercise: 


Problem:T and B lymphocytes 


a. are polymorphonuclear 

b. are involved with specific immune function 
c. proliferate excessively in leukopenia 

d. are most active against parasitic worms 


Solution: 


B 


Exercise: 


Problem: 


A patient has been experiencing severe, persistent allergy symptoms 
that are reduced when she takes an antihistamine. Before the treatment, 
this patient was likely to have had increased activity of which 
leukocyte? 


a. basophils 

b. neutrophils 

c. monocytes 

d. natural killer cells 


Solution: 
A 
Exercise: 

Problem:Thrombocytes are more accurately called 
a. clotting factors 
b. megakaryoblasts 
c. megakaryocytes 
d. platelets 

Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: 


One of the more common adverse effects of cancer chemotherapy is 
the destruction of leukocytes. Before his next scheduled chemotherapy 
treatment, a patient undergoes a blood test called an absolute 
neutrophil count (ANC), which reveals that his neutrophil count is 
1900 cells per microliter. Would his healthcare team be likely to 
proceed with his chemotherapy treatment? Why? 


Solution: 


A neutrophil count below 1800 cells per microliter is considered 
abnormal. Thus, this patient’s ANC is at the low end of the normal 
range and there would be no reason to delay chemotherapy. In clinical 
practice, most patients are given chemotherapy if their ANC is above 
1000. 


Exercise: 


Problem: 


A patient was admitted to the burn unit the previous evening suffering 
from a severe burn involving his left upper extremity and shoulder. A 
blood test reveals that he is experiencing leukocytosis. Why is this an 
expected finding? 


Solution: 


Any severe stress can increase the leukocyte count, resulting in 
leukocytosis. A burn is especially likely to increase the proliferation of 
leukocytes in order to ward off infection, a significant risk when the 
barrier function of the skin is destroyed. 


Glossary 
agranular leukocytes 


leukocytes with few granules in their cytoplasm; specifically, 
monocytes, lymphocytes, and NK cells 


B lymphocytes 
(also, B cells) lymphocytes that defend the body against specific 
pathogens and thereby provide specific immunity 


basophils 
granulocytes that stain with a basic (alkaline) stain and store histamine 
and heparin 


defensins 
antimicrobial proteins released from neutrophils and macrophages that 
create openings in the plasma membranes to kill cells 


diapedesis 
(also, emigration) process by which leukocytes squeeze through 
adjacent cells in a blood vessel wall to enter tissues 


emigration 
(also, diapedesis) process by which leukocytes squeeze through 
adjacent cells in a blood vessel wall to enter tissues 


eosinophils 
granulocytes that stain with eosin; they release antihistamines and are 
especially active against parasitic worms 


granular leukocytes 
leukocytes with abundant granules in their cytoplasm; specifically, 
neutrophils, eosinophils, and basophils 


leukemia 
cancer involving leukocytes 


leukocyte 
(also, white blood cell) colorless, nucleated blood cell, the chief 
function of which is to protect the body from disease 


leukocytosis 
excessive leukocyte proliferation 


leukopenia 
below-normal production of leukocytes 


lymphocytes 
agranular leukocytes of the lymphoid stem cell line, many of which 
function in specific immunity 


lymphoma 
form of cancer in which masses of malignant T and/or B lymphocytes 
collect in lymph nodes, the spleen, the liver, and other tissues 


lysozyme 
digestive enzyme with bactericidal properties 


megakaryocyte 
bone marrow cell that produces platelets 


memory cell 
type of B or T lymphocyte that forms after exposure to a pathogen 


monocytes 
agranular leukocytes of the myeloid stem cell line that circulate in the 
bloodstream; tissue monocytes are macrophages 


natural killer (NK) cells 
cytotoxic lymphocytes capable of recognizing cells that do not express 
“self” proteins on their plasma membrane or that contain foreign or 
abnormal markers; provide generalized, nonspecific immunity 


neutrophils 
granulocytes that stain with a neutral dye and are the most numerous 
of the leukocytes; especially active against bacteria 


polymorphonuclear 
having a lobed nucleus, as seen in some leukocytes 


positive chemotaxis 


process in which a cell is attracted to move in the direction of chemical 
stimuli 


T lymphocytes 
(also, T cells) lymphocytes that provide cellular-level immunity by 
physically attacking foreign or diseased cells 


thrombocytes 
platelets, one of the formed elements of blood that consists of cell 
fragments broken off from megakaryocytes 


thrombocytopenia 
condition in which there are too few platelets, resulting in abnormal 
bleeding (hemophilia) 


thrombocytosis 
condition in which there are too many platelets, resulting in abnormal 
clotting (thrombosis) 


Basic Structure and Function of the Nervous System 
By the end of this section, you will be able to: 


e Identify the anatomical and functional divisions of the nervous system 

¢ Relate the functional and structural differences between gray matter 
and white matter structures of the nervous system to the structure of 
neurons 

e List the basic functions of the nervous system 


The picture you have in your mind of the nervous system probably includes 
the brain, the nervous tissue contained within the cranium, and the spinal 
cord, the extension of nervous tissue within the vertebral column. That 
suggests it is made of two organs—and you may not even think of the 
spinal cord as an organ—but the nervous system is a very complex 
structure. Within the brain, many different and separate regions are 
responsible for many different and separate functions. It is as if the nervous 
system is composed of many organs that all look similar and can only be 
differentiated using tools such as the microscope or electrophysiology. In 
comparison, it is easy to see that the stomach is different than the esophagus 
or the liver, so you can imagine the digestive system as a collection of 
specific organs. 


The Central and Peripheral Nervous Systems 


The nervous system can be divided into two major regions: the central and 
peripheral nervous systems. The central nervous system (CNS) is the 
brain and spinal cord, and the peripheral nervous system (PNS) is 
everything else ([link]). The brain is contained within the cranial cavity of 
the skull, and the spinal cord is contained within the vertebral cavity of the 
vertebral column. It is a bit of an oversimplification to say that the CNS is 
what is inside these two cavities and the peripheral nervous system is 
outside of them, but that is one way to start to think about it. In actuality, 
there are some elements of the peripheral nervous system that are within the 
cranial or vertebral cavities. The peripheral nervous system is so named 
because it is on the periphery—meaning beyond the brain and spinal cord. 
Depending on different aspects of the nervous system, the dividing line 
between central and peripheral is not necessarily universal. 


Central and Peripheral Nervous System 


Central Nervous System 


Brain 


Spinal cord 


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The structures of the PNS are referred to 
as ganglia and nerves, which can be seen 
as distinct structures. The equivalent 
structures in the CNS are not obvious 
from this overall perspective and are best 
examined in prepared tissue under the 
microscope. 


Nervous tissue, present in both the CNS and PNS, contains two basic types 
of cells: neurons and glial cells. A glial cell is one of a variety of cells that 
provide a framework of tissue that supports the neurons and their activities. 
The neuron is the more functionally important of the two, in terms of the 
communicative function of the nervous system. To describe the functional 
divisions of the nervous system, it is important to understand the structure 
of aneuron. Neurons are cells and therefore have a soma, or cell body, but 
they also have extensions of the cell; each extension is generally referred to 
as a process. There is one important process that every neuron has called an 
axon, which is the fiber that connects a neuron with its target. Another type 


of process that branches off from the soma is the dendrite. Dendrites are 
responsible for receiving most of the input from other neurons. Looking at 
nervous tissue, there are regions that predominantly contain cell bodies and 
regions that are largely composed of just axons. These two regions within 
nervous system structures are often referred to as gray matter (the regions 
with many cell bodies and dendrites) or white matter (the regions with 
many axons). [link] demonstrates the appearance of these regions in the 
brain and spinal cord. The colors ascribed to these regions are what would 
be seen in “fresh,” or unstained, nervous tissue. Gray matter is not 
necessarily gray. It can be pinkish because of blood content, or even slightly 
tan, depending on how long the tissue has been preserved. But white matter 
is white because axons are insulated by a lipid-rich substance called 
myelin. Lipids can appear as white (“fatty”) material, much like the fat on a 
raw piece of chicken or beef. Actually, gray matter may have that color 
ascribed to it because next to the white matter, it is just darker—hence, 


gray. 


The distinction between gray matter and white matter is most often applied 
to central nervous tissue, which has large regions that can be seen with the 
unaided eye. When looking at peripheral structures, often a microscope is 
used and the tissue is stained with artificial colors. That is not to say that 
central nervous tissue cannot be stained and viewed under a microscope, 
but unstained tissue is most likely from the CNS—for example, a frontal 
section of the brain or cross section of the spinal cord. 

Gray Matter and White Matter 


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A brain removed during an autopsy, with 
a partial section removed, shows white 
matter surrounded by gray matter. Gray 
matter makes up the outer cortex of the 
brain. (credit: modification of work by 

“Suseno”/Wikimedia Commons) 


Regardless of the appearance of stained or unstained tissue, the cell bodies 
of neurons or axons can be located in discrete anatomical structures that 
need to be named. Those names are specific to whether the structure is 
central or peripheral. A localized collection of neuron cell bodies in the 
CNS is referred to as a nucleus. In the PNS, a cluster of neuron cell bodies 
is referred to as a ganglion. [link] indicates how the term nucleus has a few 
different meanings within anatomy and physiology. It is the center of an 
atom, where protons and neutrons are found; it is the center of a cell, where 
the DNA is found; and it is a center of some function in the CNS. There is 
also a potentially confusing use of the word ganglion (plural = ganglia) that 
has a historical explanation. In the central nervous system, there is a group 
of nuclei that are connected together and were once called the basal ganglia 
before “ganglion” became accepted as a description for a peripheral 
structure. Some sources refer to this group of nuclei as the “basal nuclei” to 
avoid confusion. 

What Is a Nucleus? 


Nucleus 


cee 


Helium atom 


(®) Proton 
O Neutron 


Electron cloud 


(a) Nucleus of an atom (b) Nucleus of a cell (c) Nucleus in the brain 


(a) The nucleus of an atom contains its 


protons and neutrons. (b) The nucleus of a 
cell is the organelle that contains DNA. (c) 
A nucleus in the CNS is a localized center 
of function with the cell bodies of several 
neurons, shown here circled in red. (credit 
c: “Was a bee”/Wikimedia Commons) 


Terminology applied to bundles of axons also differs depending on location. 
A bundle of axons, or fibers, found in the CNS is called a tract whereas the 
same thing in the PNS would be called a nerve. There is an important point 
to make about these terms, which is that they can both be used to refer to 
the same bundle of axons. When those axons are in the PNS, the term is 
nerve, but if they are CNS, the term is tract. The most obvious example of 
this is the axons that project from the retina into the brain. Those axons are 
called the optic nerve as they leave the eye, but when they are inside the 
cranium, they are referred to as the optic tract. There is a specific place 
where the name changes, which is the optic chiasm, but they are still the 
same axons ({link]). A similar situation outside of science can be described 
for some roads. Imagine a road called “Broad Street” in a town called 
“Anyville.” The road leaves Anyville and goes to the next town over, called 
“Hometown.” When the road crosses the line between the two towns and is 
in Hometown, its name changes to “Main Street.” That is the idea behind 
the naming of the retinal axons. In the PNS, they are called the optic nerve, 
and in the CNS, they are the optic tract. [link] helps to clarify which of 
these terms apply to the central or peripheral nervous systems. 

Optic Nerve Versus Optic Tract 


Left eye 


Optic nerve 
Optic chiasma 


Optic tract 
Thalamus 


Midbrain 


Occipital 
lobes 


This drawing of the connections of the 
eye to the brain shows the optic nerve 
extending from the eye to the chiasm, 
where the structure continues as the optic 
tract. The same axons extend from the eye 
to the brain through these two bundles of 
fibers, but the chiasm represents the 
border between peripheral and central. 


Note: 


ree pri 
wees OPenstax COLLEGE” 


In 2003, the Nobel Prize in Physiology or Medicine was awarded to Paul 
C. Lauterbur and Sir Peter Mansfield for discoveries related to magnetic 


resonance imaging (MRI). This is a tool to see the structures of the body 
(not just the nervous system) that depends on magnetic fields associated 
with certain atomic nuclei. The utility of this technique in the nervous 
system is that fat tissue and water appear as different shades between black 
and white. Because white matter is fatty (from myelin) and gray matter is 
not, they can be easily distinguished in MRI images. Visit the Nobel Prize 
web site to play an interactive game that demonstrates the use of this 
technology and compares it with other types of imaging technologies. 
Also, the results from an MRI session are compared with images obtained 
from X-ray or computed tomography. How do the imaging techniques 
shown in this game indicate the separation of white and gray matter 
compared with the freshly dissected tissue shown earlier? 


Structures of the CNS and PNS 


CNS PNS 
Group of Neuron Cell Bodies (i.e., gray Mncleds Ganglion 
matter) 
Bundle of Axons (i.e., white matter) Tract Nerve 


Functional Divisions of the Nervous System 


The nervous system can also be divided on the basis of its functions, but 
anatomical divisions and functional divisions are different. The CNS and 
the PNS both contribute to the same functions, but those functions can be 
attributed to different regions of the brain (such as the cerebral cortex or the 
hypothalamus) or to different ganglia in the periphery. The problem with 
trying to fit functional differences into anatomical divisions is that 
sometimes the same structure can be part of several functions. For example, 


the optic nerve carries signals from the retina that are either used for the 
conscious perception of visual stimuli, which takes place in the cerebral 
cortex, or for the reflexive responses of smooth muscle tissue that are 
processed through the hypothalamus. 


There are two ways to consider how the nervous system is divided 
functionally. First, the basic functions of the nervous system are sensation, 
integration, and response. Secondly, control of the body can be somatic or 
autonomic—divisions that are largely defined by the structures that are 
involved in the response. There is also a region of the peripheral nervous 
system that is called the enteric nervous system that is responsible for a 
specific set of the functions within the realm of autonomic control related to 
gastrointestinal functions. 


Basic Functions 


The nervous system is involved in receiving information about the 
environment around us (sensation) and generating responses to that 
information (motor responses). The nervous system can be divided into 
regions that are responsible for sensation (sensory functions) and for the 
response (motor functions). But there is a third function that needs to be 
included. Sensory input needs to be integrated with other sensations, as well 
as with memories, emotional state, or learning (cognition). Some regions of 
the nervous system are termed integration or association areas. The process 
of integration combines sensory perceptions and higher cognitive functions 
such as memories, learning, and emotion to produce a response. 


Sensation. The first major function of the nervous system is sensation— 
receiving information about the environment to gain input about what is 
happening outside the body (or, sometimes, within the body). The sensory 
functions of the nervous system register the presence of a change from 
homeostasis or a particular event in the environment, known as a stimulus. 
The senses we think of most are the “big five”: taste, smell, touch, sight, 
and hearing. The stimuli for taste and smell are both chemical substances 
(molecules, compounds, ions, etc.), touch is physical or mechanical stimuli 
that interact with the skin, sight is light stimuli, and hearing is the 


perception of sound, which is a physical stimulus similar to some aspects of 
touch. There are actually more senses than just those, but that list represents 
the major senses. Those five are all senses that receive stimuli from the 
outside world, and of which there is conscious perception. Additional 
sensory stimuli might be from the internal environment (inside the body), 
such as the stretch of an organ wall or the concentration of certain ions in 
the blood. 


Response. The nervous system produces a response on the basis of the 
stimuli perceived by sensory structures. An obvious response would be the 
movement of muscles, such as withdrawing a hand from a hot stove, but 
there are broader uses of the term. The nervous system can cause the 
contraction of all three types of muscle tissue. For example, skeletal muscle 
contracts to move the skeleton, cardiac muscle is influenced as heart rate 
increases during exercise, and smooth muscle contracts as the digestive 
system moves food along the digestive tract. Responses also include the 
neural control of glands in the body as well, such as the production and 
secretion of sweat by the eccrine and merocrine sweat glands found in the 
skin to lower body temperature. 


Responses can be divided into those that are voluntary or conscious 
(contraction of skeletal muscle) and those that are involuntary (contraction 
of smooth muscles, regulation of cardiac muscle, activation of glands). 
Voluntary responses are governed by the somatic nervous system and 
involuntary responses are governed by the autonomic nervous system, 
which are discussed in the next section. 


Integration. Stimuli that are received by sensory structures are 
communicated to the nervous system where that information is processed. 
This is called integration. Stimuli are compared with, or integrated with, 
other stimuli, memories of previous stimuli, or the state of a person at a 
particular time. This leads to the specific response that will be generated. 
Seeing a baseball pitched to a batter will not automatically cause the batter 
to swing. The trajectory of the ball and its speed will need to be considered. 
Maybe the count is three balls and one strike, and the batter wants to let this 
pitch go by in the hope of getting a walk to first base. Or maybe the batter’s 
team is so far ahead, it would be fun to just swing away. 


Controlling the Body 


The nervous system can be divided into two parts mostly on the basis of a 
functional difference in responses. The somatic nervous system (SNS) is 
responsible for conscious perception and voluntary motor responses. 
Voluntary motor response means the contraction of skeletal muscle, but 
those contractions are not always voluntary in the sense that you have to 
want to perform them. Some somatic motor responses are reflexes, and 
often happen without a conscious decision to perform them. If your friend 
jumps out from behind a corner and yells “Boo!” you will be startled and 
you might scream or leap back. You didn’t decide to do that, and you may 
not have wanted to give your friend a reason to laugh at your expense, but it 
is areflex involving skeletal muscle contractions. Other motor responses 
become automatic (in other words, unconscious) as a person learns motor 
skills (referred to as “habit learning” or “procedural memory”). 


The autonomic nervous system (ANS) is responsible for involuntary 
control of the body, usually for the sake of homeostasis (regulation of the 
internal environment). Sensory input for autonomic functions can be from 
sensory structures tuned to external or internal environmental stimuli. The 
motor output extends to smooth and cardiac muscle as well as glandular 
tissue. The role of the autonomic system is to regulate the organ systems of 
the body, which usually means to control homeostasis. Sweat glands, for 
example, are controlled by the autonomic system. When you are hot, 
sweating helps cool your body down. That is a homeostatic mechanism. But 
when you are nervous, you might start sweating also. That is not 
homeostatic, it is the physiological response to an emotional state. 


There is another division of the nervous system that describes functional 
responses. The enteric nervous system (ENS) is responsible for 
controlling the smooth muscle and glandular tissue in your digestive 
system. It is a large part of the PNS, and is not dependent on the CNS. It is 
sometimes valid, however, to consider the enteric system to be a part of the 
autonomic system because the neural structures that make up the enteric 
system are a component of the autonomic output that regulates digestion. 
There are some differences between the two, but for our purposes here there 


will be a good bit of overlap. See [link] for examples of where these 


divisions of the nervous system can be found. 
Somatic, Autonomic, and Enteric Structures of the Nervous System 


Brain (CNS) 
Perception and processing of sensory 
stimuli (somatic/autonomic) 
Execution of voluntary motor 
responses (somatic) 
Regulation of homeostatic 
mechanisms (autonomic) 


Spinal cord (CNS) 

Initiation of reflexes from ventral 
horn (somatic) and lateral horn 
(autonomic) gray matter 

Pathways for sensory and motor 
functions between periphery 
and brain (somatic/autonomic) 


Nerves (PNS) 
Fibers of sensory and motor 
neurons (somatic/autonomic) 


Ganglia (PNS) 

Reception of sensory stimuli by 
dorsal root and cranial ganglia 
(somatic/autonomic) 

Relay of visceral motor responses 

Digestive tract (ENS) by autonomic ganglia (autonomic) 

The enteric nervous system | 

(ENS), located in the digestive 

tract, is responsible for autonomous 

functions and can operate independently 

of the brain and spinal cord. 


Somatic structures include the spinal nerves, both motor and 
sensory fibers, as well as the sensory ganglia (posterior root 
ganglia and cranial nerve ganglia). Autonomic structures are found 
in the nerves also, but include the sympathetic and parasympathetic 
ganglia. The enteric nervous system includes the nervous tissue 
within the organs of the digestive tract. 


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—o 


Visit this site to read about a woman that notices that her daughter is 
having trouble walking up the stairs. This leads to the discovery of a 
hereditary condition that affects the brain and spinal cord. The 
electromyography and MRI tests indicated deficiencies in the spinal cord 
and cerebellum, both of which are responsible for controlling coordinated 
movements. To what functional division of the nervous system would these 
structures belong? 


Note: 

Everyday Connection 

How Much of Your Brain Do You Use? 

Have you ever heard the claim that humans only use 10 percent of their 
brains? Maybe you have seen an advertisement on a website saying that 
there is a secret to unlocking the full potential of your mind—as if there 
were 90 percent of your brain sitting idle, just waiting for you to use it. If 
you see an ad like that, don’t click. It isn’t true. 

An easy way to see how much of the brain a person uses is to take 
measurements of brain activity while performing a task. An example of 
this kind of measurement is functional magnetic resonance imaging 
(fMRI), which generates a map of the most active areas and can be 
generated and presented in three dimensions ([link]). This procedure is 
different from the standard MRI technique because it is measuring changes 
in the tissue in time with an experimental condition or event. 

fMRI 


This {MRI shows activation of the visual 
cortex in response to visual stimuli. 
(credit: “Superborsuk”/Wikimedia 
Commons) 


The underlying assumption is that active nervous tissue will have greater 
blood flow. By having the subject perform a visual task, activity all over 
the brain can be measured. Consider this possible experiment: the subject 
is told to look at a screen with a black dot in the middle (a fixation point). 
A photograph of a face is projected on the screen away from the center. 
The subject has to look at the photograph and decipher what it is. The 
subject has been instructed to push a button if the photograph is of 
someone they recognize. The photograph might be of a celebrity, so the 
subject would press the button, or it might be of a random person unknown 
to the subject, so the subject would not press the button. 

In this task, visual sensory areas would be active, integrating areas would 
be active, motor areas responsible for moving the eyes would be active, 
and motor areas for pressing the button with a finger would be active. 
Those areas are distributed all around the brain and the [MRI images 
would show activity in more than just 10 percent of the brain (some 
evidence suggests that about 80 percent of the brain is using energy— 


based on blood flow to the tissue—during well-defined tasks similar to the 
one suggested above). This task does not even include all of the functions 
the brain performs. There is no language response, the body is mostly lying 
still in the MRI machine, and it does not consider the autonomic functions 
that would be ongoing in the background. 


Chapter Review 


The nervous system can be separated into divisions on the basis of anatomy 
and physiology. The anatomical divisions are the central and peripheral 
nervous systems. The CNS is the brain and spinal cord. The PNS is 
everything else. Functionally, the nervous system can be divided into those 
regions that are responsible for sensation, those that are responsible for 
integration, and those that are responsible for generating responses. All of 
these functional areas are found in both the central and peripheral anatomy. 


Considering the anatomical regions of the nervous system, there are specific 
names for the structures within each division. A localized collection of 
neuron cell bodies is referred to as a nucleus in the CNS and as a ganglion 
in the PNS. A bundle of axons is referred to as a tract in the CNS and as a 
nerve in the PNS. Whereas nuclei and ganglia are specifically in the central 
or peripheral divisions, axons can cross the boundary between the two. A 
single axon can be part of a nerve and a tract. The name for that specific 
structure depends on its location. 


Nervous tissue can also be described as gray matter and white matter on the 
basis of its appearance in unstained tissue. These descriptions are more 
often used in the CNS. Gray matter is where nuclei are found and white 
matter is where tracts are found. In the PNS, ganglia are basically gray 
matter and nerves are white matter. 


The nervous system can also be divided on the basis of how it controls the 
body. The somatic nervous system (SNS) is responsible for functions that 
result in moving skeletal muscles. Any sensory or integrative functions that 
result in the movement of skeletal muscle would be considered somatic. 
The autonomic nervous system (ANS) is responsible for functions that 


affect cardiac or smooth muscle tissue, or that cause glands to produce their 
secretions. Autonomic functions are distributed between central and 
peripheral regions of the nervous system. The sensations that lead to 
autonomic functions can be the same sensations that are part of initiating 
somatic responses. Somatic and autonomic integrative functions may 
overlap as well. 


A special division of the nervous system is the enteric nervous system, 
which is responsible for controlling the digestive organs. Parts of the 
autonomic nervous system overlap with the enteric nervous system. The 
enteric nervous system is exclusively found in the periphery because it is 
the nervous tissue in the organs of the digestive system. 


Interactive Link Questions 


Exercise: 


Problem: 


In 2003, the Nobel Prize in Physiology or Medicine was awarded to 
Paul C. Lauterbur and Sir Peter Mansfield for discoveries related to 
magnetic resonance imaging (MRI). This is a tool to see the structures 
of the body (not just the nervous system) that depends on magnetic 
fields associated with certain atomic nuclei. The utility of this 
technique in the nervous system is that fat tissue and water appear as 
different shades between black and white. Because white matter is 
fatty (from myelin) and gray matter is not, they can be easily 
distinguished in MRI images. Visit the Nobel Prize website to play an 
interactive game that demonstrates the use of this technology and 
compares it with other types of imaging technologies. Also, the results 
from an MRI session are compared with images obtained from x-ray or 
computed tomography. How do the imaging techniques shown in this 
game indicate the separation of white and gray matter compared with 
the freshly dissected tissue shown earlier? 


Solution: 


MRI uses the relative amount of water in tissue to distinguish different 
areas, SO gray and white matter in the nervous system can be seen 
clearly in these images. 


Exercise: 
Problem: 
Visit this site to read about a woman that notices that her daughter is 
having trouble walking up the stairs. This leads to the discovery of a 
hereditary condition that affects the brain and spinal cord. The 
electromyography and MRI tests indicated deficiencies in the spinal 
cord and cerebellum, both of which are responsible for controlling 


coordinated movements. To what functional division of the nervous 
system would these structures belong? 


Solution: 

They are part of the somatic nervous system, which is responsible for 

voluntary movements such as walking or climbing the stairs. 
Review Questions 


Exercise: 


Problem: 


Which of the following cavities contains a component of the central 
nervous system? 


a. abdominal 
b. pelvic 

c. cranial 

d. thoracic 


Solution: 


C 


Exercise: 


Problem: 
Which structure predominates in the white matter of the brain? 


a. myelinated axons 

b. neuronal cell bodies 

c. ganglia of the parasympathetic nerves 

d. bundles of dendrites from the enteric nervous system 


Solution: 


A 
Exercise: 


Problem: 


Which part of a neuron transmits an electrical signal to a target cell? 


a. dendrites 
b. soma 
c. cell body 
d. axon 


Solution: 


D 
Exercise: 


Problem: 


Which term describes a bundle of axons in the peripheral nervous 
system? 


a. nucleus 
b. ganglion 


c. tract 
d. nerve 


Solution: 


D 
Exercise: 


Problem: 


Which functional division of the nervous system would be responsible 
for the physiological changes seen during exercise (e.g., increased 
heart rate and sweating)? 


a. Somatic 

b. autonomic 
c. enteric 

d. central 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


What responses are generated by the nervous system when you run on 
a treadmill? Include an example of each type of tissue that is under 
nervous system control. 


Solution: 


Running on a treadmill involves contraction of the skeletal muscles in 
the legs, increase in contraction of the cardiac muscle of the heart, and 
the production and secretion of sweat in the skin to stay cool. 


Exercise: 
Problem: 


When eating food, what anatomical and functional divisions of the 
nervous system are involved in the perceptual experience? 


Solution: 


The sensation of taste associated with eating is sensed by nerves in the 
periphery that are involved in sensory and somatic functions. 


References 


Kramer, PD. Listening to prozac. 1st ed. New York (NY): Penguin Books; 
1993: 


Glossary 


autonomic nervous system (ANS) 
functional division of the nervous system that is responsible for 
homeostatic reflexes that coordinate control of cardiac and smooth 
muscle, as well as glandular tissue 


axon 
single process of the neuron that carries an electrical signal (action 
potential) away from the cell body toward a target cell 


brain 
the large organ of the central nervous system composed of white and 
gray matter, contained within the cranium and continuous with the 
spinal cord 


central nervous system (CNS) 


anatomical division of the nervous system located within the cranial 
and vertebral cavities, namely the brain and spinal cord 


dendrite 
one of many branchlike processes that extends from the neuron cell 
body and functions as a contact for incoming signals (synapses) from 
other neurons or sensory cells 


enteric nervous system (ENS) 
neural tissue associated with the digestive system that is responsible 
for nervous control through autonomic connections 


ganglion 
localized collection of neuron cell bodies in the peripheral nervous 
system 


glial cell 
one of the various types of neural tissue cells responsible for 
maintenance of the tissue, and largely responsible for supporting 
neurons 


gray matter 
regions of the nervous system containing cell bodies of neurons with 
few or no myelinated axons; actually may be more pink or tan in color, 
but called gray in contrast to white matter 


integration 
nervous system function that combines sensory perceptions and higher 
cognitive functions (memories, learning, emotion, etc.) to produce a 
response 


myelin 
lipid-rich insulating substance surrounding the axons of many neurons, 
allowing for faster transmission of electrical signals 


nerve 
cord-like bundle of axons located in the peripheral nervous system that 
transmits sensory input and response output to and from the central 


nervous system 


neuron 
neural tissue cell that is primarily responsible for generating and 
propagating electrical signals into, within, and out of the nervous 
system 


nucleus 
in the nervous system, a localized collection of neuron cell bodies that 
are functionally related; a “center” of neural function 


peripheral nervous system (PNS) 
anatomical division of the nervous system that is largely outside the 
cranial and vertebral cavities, namely all parts except the brain and 
spinal cord 


process 
in cells, an extension of a cell body; in the case of neurons, this 
includes the axon and dendrites 


response 
nervous system function that causes a target tissue (muscle or gland) to 
produce an event as a consequence to stimuli 


sensation 
nervous system function that receives information from the 
environment and translates it into the electrical signals of nervous 
tissue 


soma 
in neurons, that portion of the cell that contains the nucleus; the cell 
body, as opposed to the cell processes (axons and dendrites) 


somatic nervous system (SNS) 
functional division of the nervous system that is concerned with 
conscious perception, voluntary movement, and skeletal muscle 
reflexes 


spinal cord 
organ of the central nervous system found within the vertebral cavity 
and connected with the periphery through spinal nerves; mediates 
reflex behaviors 


stimulus 
an event in the external or internal environment that registers as 
activity in a sensory neuron 


tract 
bundle of axons in the central nervous system having the same 
function and point of origin 


white matter 
regions of the nervous system containing mostly myelinated axons, 
making the tissue appear white because of the high lipid content of 
myelin 


Nervous Tissue 
By the end of this section, you will be able to: 


e Describe the basic structure of a neuron 

e Identify the different types of neurons on the basis of polarity 
e List the glial cells of the CNS and describe their function 
List the glial cells of the PNS and describe their function 


Nervous tissue is composed of two types of cells, neurons and glial cells. 
Neurons are the primary type of cell that most anyone associates with the 
nervous system. They are responsible for the computation and 
communication that the nervous system provides. They are electrically 
active and release chemical signals to target cells. Glial cells, or glia, are 
known to play a supporting role for nervous tissue. Ongoing research 
pursues an expanded role that glial cells might play in signaling, but 
neurons are still considered the basis of this function. Neurons are 
important, but without glial support they would not be able to perform their 
function. 


Neurons 


Neurons are the cells considered to be the basis of nervous tissue. They are 
responsible for the electrical signals that communicate information about 
sensations, and that produce movements in response to those stimuli, along 
with inducing thought processes within the brain. An important part of the 
function of neurons is in their structure, or shape. The three-dimensional 
shape of these cells makes the immense numbers of connections within the 
nervous system possible. 


Parts of a Neuron 


As you leamed in the first section, the main part of a neuron is the cell 
body, which is also known as the soma (soma = “body”). The cell body 
contains the nucleus and most of the major organelles. But what makes 
neurons special is that they have many extensions of their cell membranes, 
which are generally referred to as processes. Neurons are usually described 


as having one, and only one, axon—a fiber that emerges from the cell body 
and projects to target cells. That single axon can branch repeatedly to 
communicate with many target cells. It is the axon that propagates the nerve 
impulse, which is communicated to one or more cells. The other processes 
of the neuron are dendrites, which receive information from other neurons 
at specialized areas of contact called synapses. The dendrites are usually 
highly branched processes, providing locations for other neurons to 
communicate with the cell body. Information flows through a neuron from 
the dendrites, across the cell body, and down the axon. This gives the 
neuron a polarity—meaning that information flows in this one direction. 
[link] shows the relationship of these parts to one another. 

Parts of a Neuron 


Cell body (soma) 


Axon Oligodendrocyte 


Cell membrane 


Dendrites 


Node of Ranvier 


Myelin sheath 


Synapse 


The major parts of the neuron are labeled 
on a multipolar neuron from the CNS. 


Where the axon emerges from the cell body, there is a special region 
referred to as the axon hillock. This is a tapering of the cell body toward 
the axon fiber. Within the axon hillock, the cytoplasm changes to a solution 
of limited components called axoplasm. Because the axon hillock 
represents the beginning of the axon, it is also referred to as the initial 
segment. 


Many axons are wrapped by an insulating substance called myelin, which is 
actually made from glial cells. Myelin acts as insulation much like the 
plastic or rubber that is used to insulate electrical wires. A key difference 
between myelin and the insulation on a wire is that there are gaps in the 
myelin covering of an axon. Each gap is called a node of Ranvier and is 
important to the way that electrical signals travel down the axon. The length 
of the axon between each gap, which is wrapped in myelin, is referred to as 
an axon segment. At the end of the axon is the axon terminal, where there 
are usually several branches extending toward the target cell, each of which 
ends in an enlargement called a synaptic end bulb. These bulbs are what 
make the connection with the target cell at the synapse. 


Note: 

ae COLLEGE 
FRR eT 
" Lal 
[a] ate ce 


Visit this site to learn about how nervous tissue is composed of neurons 
and glial cells. Neurons are dynamic cells with the ability to make a vast 
number of connections, to respond incredibly quickly to stimuli, and to 
initiate movements on the basis of those stimuli. They are the focus of 
intense research because failures in physiology can lead to devastating 
illnesses. Why are neurons only found in animals? Based on what this 
article says about neuron function, why wouldn't they be helpful for plants 
or microorganisms? 


Types of Neurons 


There are many neurons in the nervous system—a number in the trillions. 
And there are many different types of neurons. They can be classified by 
many different criteria. The first way to classify them is by the number of 


processes attached to the cell body. Using the standard model of neurons, 
one of these processes is the axon, and the rest are dendrites. Because 
information flows through the neuron from dendrites or cell bodies toward 
the axon, these names are based on the neuron's polarity ([link]). 

Neuron Classification by Shape 


Bipolar neuron 


Unipolar neuron 


Multipolar neuron 


Unipolar cells have one process that includes 
both the axon and dendrite. Bipolar cells have 
two processes, the axon and a dendrite. 
Multipolar cells have more than two processes, 
the axon and two or more dendrites. 


Unipolar cells have only one process emerging from the cell. True unipolar 
cells are only found in invertebrate animals, so the unipolar cells in humans 
are more appropriately called “pseudo-unipolar” cells. Invertebrate unipolar 
cells do not have dendrites. Human unipolar cells have an axon that 
emerges from the cell body, but it splits so that the axon can extend along a 
very long distance. At one end of the axon are dendrites, and at the other 
end, the axon forms synaptic connections with a target. Unipolar cells are 
exclusively sensory neurons and have two unique characteristics. First, their 
dendrites are receiving sensory information, sometimes directly from the 
stimulus itself. Secondly, the cell bodies of unipolar neurons are always 
found in ganglia. Sensory reception is a peripheral function (those dendrites 


are in the periphery, perhaps in the skin) so the cell body is in the periphery, 
though closer to the CNS in a ganglion. The axon projects from the dendrite 
endings, past the cell body in a ganglion, and into the central nervous 
system. 


Bipolar cells have two processes, which extend from each end of the cell 
body, opposite to each other. One is the axon and one the dendrite. Bipolar 
cells are not very common. They are found mainly in the olfactory 
epithelium (where smell stimuli are sensed), and as part of the retina. 


Multipolar neurons are all of the neurons that are not unipolar or bipolar. 
They have one axon and two or more dendrites (usually many more). With 
the exception of the unipolar sensory ganglion cells, and the two specific 
bipolar cells mentioned above, all other neurons are multipolar. Some 
cutting edge research suggests that certain neurons in the CNS do not 
conform to the standard model of “one, and only one” axon. Some sources 
describe a fourth type of neuron, called an anaxonic neuron. The name 
suggests that it has no axon (an- = “without”), but this is not accurate. 
Anaxonic neurons are very small, and if you look through a microscope at 
the standard resolution used in histology (approximately 400X to 1000X 
total magnification), you will not be able to distinguish any process 
specifically as an axon or a dendrite. Any of those processes can function as 
an axon depending on the conditions at any given time. Nevertheless, even 
if they cannot be easily seen, and one specific process is definitively the 
axon, these neurons have multiple processes and are therefore multipolar. 


Neurons can also be classified on the basis of where they are found, who 
found them, what they do, or even what chemicals they use to communicate 
with each other. Some neurons referred to in this section on the nervous 
system are named on the basis of those sorts of classifications ((link]). For 
example, a multipolar neuron that has a very important role to play in a part 
of the brain called the cerebellum is known as a Purkinje (commonly 
pronounced per-KIN-gee) cell. It is named after the anatomist who 
discovered it (Jan Evangilista Purkinje, 1787-1869). 

Other Neuron Classifications 


(a) Pyramidal cell of the (b) Purkinje cell of the (c) Olfactory cells in the olfactory 
cerebral cortex cerebellar cortex epithelium and olfactory bulbs 


Three examples of neurons that are classified on 
the basis of other criteria. (a) The pyramidal cell is 
a multipolar cell with a cell body that is shaped 
something like a pyramid. (b) The Purkinje cell in 
the cerebellum was named after the scientist who 
originally described it. (c) Olfactory neurons are 
named for the functional group with which they 
belong. 


Glial Cells 


Glial cells, or neuroglia or simply glia, are the other type of cell found in 
nervous tissue. They are considered to be supporting cells, and many 
functions are directed at helping neurons complete their function for 
communication. The name glia comes from the Greek word that means 
“glue,” and was coined by the German pathologist Rudolph Virchow, who 
wrote in 1856: “This connective substance, which is in the brain, the spinal 
cord, and the special sense nerves, is a kind of glue (neuroglia) in which the 
nervous elements are planted.” Today, research into nervous tissue has 
shown that there are many deeper roles that these cells play. And research 
may find much more about them in the future. 


There are six types of glial cells. Four of them are found in the CNS and 
two are found in the PNS. [link] outlines some common characteristics and 
functions. 


Glial Cell Types by Location and Basic Function 


CNS glia PNS glia Basic function 
Satellite 
Astrocyte cell Support 
Oligodendrocyte all Insulation, myelination 
; Immune surveillance and 
Microglia - ; 
phagocytosis 
Ependymal cell - Creating CSF 
Glial Cells of the CNS 


One cell providing support to neurons of the CNS is the astrocyte, so 
named because it appears to be star-shaped under the microscope (astro- = 
“star”). Astrocytes have many processes extending from their main cell 
body (not axons or dendrites like neurons, just cell extensions). Those 
processes extend to interact with neurons, blood vessels, or the connective 
tissue covering the CNS that is called the pia mater ({link]). Generally, they 
are supporting cells for the neurons in the central nervous system. Some 
ways in which they support neurons in the central nervous system are by 
maintaining the concentration of chemicals in the extracellular space, 


removing excess signaling molecules, reacting to tissue damage, and 
contributing to the blood-brain barrier (BBB). The blood-brain barrier is a 
physiological barrier that keeps many substances that circulate in the rest of 
the body from getting into the central nervous system, restricting what can 
cross from circulating blood into the CNS. Nutrient molecules, such as 
glucose or amino acids, can pass through the BBB, but other molecules 
cannot. This actually causes problems with drug delivery to the CNS. 
Pharmaceutical companies are challenged to design drugs that can cross the 
BBB as well as have an effect on the nervous system. 

Glial Cells of the CNS 


Microglial cell if aw 


Astrocyte 


7} Ependymal cells al Oligodendrocytes 
The CNS has astrocytes, oligodendrocytes, 
microglia, and ependymal cells that support the 
neurons of the CNS in several ways. 


Like a few other parts of the body, the brain has a privileged blood supply. 
Very little can pass through by diffusion. Most substances that cross the 
wall of a blood vessel into the CNS must do so through an active transport 
process. Because of this, only specific types of molecules can enter the 
CNS. Glucose—the primary energy source—is allowed, as are amino acids. 
Water and some other small particles, like gases and ions, can enter. But 


most everything else cannot, including white blood cells, which are one of 
the body’s main lines of defense. While this barrier protects the CNS from 
exposure to toxic or pathogenic substances, it also keeps out the cells that 
could protect the brain and spinal cord from disease and damage. The BBB 
also makes it harder for pharmaceuticals to be developed that can affect the 
nervous system. Aside from finding efficacious substances, the means of 
delivery is also crucial. 


Also found in CNS tissue is the oligodendrocyte, sometimes called just 
“oligo,” which is the glial cell type that insulates axons in the CNS. The 
name means “cell of a few branches” (oligo- = “few”; dendro- = 
“branches”; -cyte = “cell”). There are a few processes that extend from the 
cell body. Each one reaches out and surrounds an axon to insulate it in 
myelin. One oligodendrocyte will provide the myelin for multiple axon 
segments, either for the same axon or for separate axons. The function of 
myelin will be discussed below. 


Microglia are, as the name implies, smaller than most of the other glial 
cells. Ongoing research into these cells, although not entirely conclusive, 
suggests that they may originate as white blood cells, called macrophages, 
that become part of the CNS during early development. While their origin is 
not conclusively determined, their function is related to what macrophages 
do in the rest of the body. When macrophages encounter diseased or 
damaged cells in the rest of the body, they ingest and digest those cells or 
the pathogens that cause disease. Microglia are the cells in the CNS that can 
do this in normal, healthy tissue, and they are therefore also referred to as 
CNS-resident macrophages. 


The ependymal cell is a glial cell that filters blood to make cerebrospinal 
fluid (CSF), the fluid that circulates through the CNS. Because of the 
privileged blood supply inherent in the BBB, the extracellular space in 
nervous tissue does not easily exchange components with the blood. 
Ependymal cells line each ventricle, one of four central cavities that are 
remnants of the hollow center of the neural tube formed during the 
embryonic development of the brain. The choroid plexus is a specialized 
structure in the ventricles where ependymal cells come in contact with 
blood vessels and filter and absorb components of the blood to produce 


cerebrospinal fluid. Because of this, ependymal cells can be considered a 
component of the BBB, or a place where the BBB breaks down. These glial 
cells appear similar to epithelial cells, making a single layer of cells with 
little intracellular space and tight connections between adjacent cells. They 
also have cilia on their apical surface to help move the CSF through the 
ventricular space. The relationship of these glial cells to the structure of the 
CNS is seen in [link]. 


Glial Cells of the PNS 


One of the two types of glial cells found in the PNS is the satellite cell. 
Satellite cells are found in sensory and autonomic ganglia, where they 
surround the cell bodies of neurons. This accounts for the name, based on 
their appearance under the microscope. They provide support, performing 
similar functions in the periphery as astrocytes do in the CNS—except, of 
course, for establishing the BBB. 


The second type of glial cell is the Schwann cell, which insulate axons with 
myelin in the periphery. Schwann cells are different than oligodendrocytes, 
in that a Schwann cell wraps around a portion of only one axon segment 
and no others. Oligodendrocytes have processes that reach out to multiple 
axon segments, whereas the entire Schwann cell surrounds just one axon 
segment. The nucleus and cytoplasm of the Schwann cell are on the edge of 
the myelin sheath. The relationship of these two types of glial cells to 
ganglia and nerves in the PNS is seen in [link]. 

Glial Cells of the PNS 


Peripheral ganglionic 
neuron cell body 
(unipolar cell) 


Satellite cells 
Schwann cells 


Axon 


The PNS has satellite cells and Schwann cells. 


Myelin 


The insulation for axons in the nervous system is provided by glial cells, 
oligodendrocytes in the CNS, and Schwann cells in the PNS. Whereas the 
manner in which either cell is associated with the axon segment, or 
segments, that it insulates is different, the means of myelinating an axon 
segment is mostly the same in the two situations. Myelin is a lipid-rich 
sheath that surrounds the axon and by doing so creates a myelin sheath that 
facilitates the transmission of electrical signals along the axon. The lipids 
are essentially the phospholipids of the glial cell membrane. Myelin, 
however, is more than just the membrane of the glial cell. It also includes 
important proteins that are integral to that membrane. Some of the proteins 
help to hold the layers of the glial cell membrane closely together. 


The appearance of the myelin sheath can be thought of as similar to the 
pastry wrapped around a hot dog for “pigs in a blanket” or a similar food. 
The glial cell is wrapped around the axon several times with little to no 
cytoplasm between the glial cell layers. For oligodendrocytes, the rest of the 
cell is separate from the myelin sheath as a cell process extends back 


toward the cell body. A few other processes provide the same insulation for 
other axon segments in the area. For Schwann cells, the outermost layer of 
the cell membrane contains cytoplasm and the nucleus of the cell as a bulge 
on one side of the myelin sheath. During development, the glial cell is 
loosely or incompletely wrapped around the axon ([link]a). The edges of 
this loose enclosure extend toward each other, and one end tucks under the 
other. The inner edge wraps around the axon, creating several layers, and 
the other edge closes around the outside so that the axon is completely 
enclosed. 


cs 
mess Openstax COLLEGE 


View the University of Michigan WebScope to see an electron micrograph 
of a cross-section of a myelinated nerve fiber. The axon contains 
microtubules and neurofilaments that are bounded by a plasma membrane 
known as the axolemma. Outside the plasma membrane of the axon is the 
myelin sheath, which is composed of the tightly wrapped plasma 
membrane of a Schwann cell. What aspects of the cells in this image react 
with the stain to make them a deep, dark, black color, such as the multiple 
layers that are the myelin sheath? 


Myelin sheaths can extend for one or two millimeters, depending on the 
diameter of the axon. Axon diameters can be as small as 1 to 20 
micrometers. Because a micrometer is 1/1000 of a millimeter, this means 
that the length of a myelin sheath can be 100-1000 times the diameter of 
the axon. [link], [link], and [link] show the myelin sheath surrounding an 
axon segment, but are not to scale. If the myelin sheath were drawn to scale, 


the neuron would have to be immense—possibly covering an entire wall of 
the room in which you are sitting. 


The Process of Myelination 
Nucleus Axon Node of Ranvier 


WebScopes 
Wl 15704 x 16044 size 720.85MB mag 20X 
> |Gau FP SSH d Pe Oo» s 


Myelin sheath 


External lamina 


Endonerium 
(collagen) 


(b) 


Myelinating glia wrap several layers of cell membrane around 
the cell membrane of an axon segment. A single Schwann cell 
insulates a segment of a peripheral nerve, whereas in the CNS, 
an oligodendrocyte may provide insulation for a few separate 
axon segments. EM x 1,460,000. (Micrograph provided by the 
Regents of University of Michigan Medical School © 2012) 


Note: 

Disorders of the... 

Nervous Tissue 

Several diseases can result from the demyelination of axons. The causes of 
these diseases are not the same; some have genetic causes, some are caused 
by pathogens, and others are the result of autoimmune disorders. Though 
the causes are varied, the results are largely similar. The myelin insulation 
of axons is compromised, making electrical signaling slower. 

Multiple sclerosis (MS) is one such disease. It is an example of an 
autoimmune disease. The antibodies produced by lymphocytes (a type of 
white blood cell) mark myelin as something that should not be in the body. 
This causes inflammation and the destruction of the myelin in the central 
nervous system. As the insulation around the axons is destroyed by the 
disease, scarring becomes obvious. This is where the name of the disease 
comes from; sclerosis means hardening of tissue, which is what a scar is. 
Multiple scars are found in the white matter of the brain and spinal cord. 
The symptoms of MS include both somatic and autonomic deficits. Control 
of the musculature is compromised, as is control of organs such as the 
bladder. 

Guillain-Barré (pronounced gee- YAN bah-RAY) syndrome is an example 
of a demyelinating disease of the peripheral nervous system. It is also the 
result of an autoimmune reaction, but the inflammation is in peripheral 
nerves. Sensory symptoms or motor deficits are common, and autonomic 
failures can lead to changes in the heart rhythm or a drop in blood pressure, 
especially when standing, which causes dizziness. 


Chapter Review 


Nervous tissue contains two major cell types, neurons and glial cells. 
Neurons are the cells responsible for communication through electrical 


signals. Glial cells are supporting cells, maintaining the environment around 
the neurons. 


Neurons are polarized cells, based on the flow of electrical signals along 
their membrane. Signals are received at the dendrites, are passed along the 
cell body, and propagate along the axon towards the target, which may be 
another neuron, muscle tissue, or a gland. Many axons are insulated by a 
lipid-rich substance called myelin. Specific types of glial cells provide this 
insulation. 


Several types of glial cells are found in the nervous system, and they can be 
categorized by the anatomical division in which they are found. In the CNS, 
astrocytes, oligodendrocytes, microglia, and ependymal cells are found. 
Astrocytes are important for maintaining the chemical environment around 
the neuron and are crucial for regulating the blood-brain barrier. 
Oligodendrocytes are the myelinating glia in the CNS. Microglia act as 
phagocytes and play a role in immune surveillance. Ependymal cells are 
responsible for filtering the blood to produce cerebrospinal fluid, which is a 
circulatory fluid that performs some of the functions of blood in the brain 
and spinal cord because of the BBB. In the PNS, satellite cells are 
supporting cells for the neurons, and Schwann cells insulate peripheral 
axons. 


Interactive Link Questions 


Exercise: 


Problem: 


Visit this site to learn about how nervous tissue is composed of 
neurons and glial cells. The neurons are dynamic cells with the ability 
to make a vast number of connections and to respond incredibly 
quickly to stimuli and to initiate movements based on those stimuli. 
They are the focus of intense research as failures in physiology can 
lead to devastating illnesses. Why are neurons only found in animals? 
Based on what this article says about neuron function, why wouldn’t 
they be helpful for plants or microorganisms? 


Solution: 


Neurons enable thought, perception, and movement. Plants do not 
move, so they do not need this type of tissue. Microorganisms are too 
small to have a nervous system. Many are single-celled, and therefore 
have organelles for perception and movement. 


Exercise: 


Problem: 


View the University of Michigan Webscope to see an electron 
micrograph of a cross-section of a myelinated nerve fiber. The axon 
contains microtubules and neurofilaments, bounded by a plasma 
membrane known as the axolemma. Outside the plasma membrane of 
the axon is the myelin sheath, which is composed of the tightly 
wrapped plasma membrane of a Schwann cell. What aspects of the 
cells in this image react with the stain that makes them the deep, dark, 
black color, such as the multiple layers that are the myelin sheath? 


Solution: 
Lipid membranes, such as the cell membrane and organelle 
membranes. 

Review Questions 


Exercise: 


Problem: 
What type of glial cell provides myelin for the axons in a tract? 


a. oligodendrocyte 
b. astrocyte 

c. Schwann cell 

d. satellite cell 


Solution: 


A 


Exercise: 


Problem: Which part of a neuron contains the nucleus? 


a. dendrite 

b. soma 

c. axon 

d. synaptic end bulb 


Solution: 


B 
Exercise: 


Problem: 


Which of the following substances is least able to cross the blood-brain 
barrier? 


a. water 

b. sodium ions 

c. glucose 

d. white blood cells 


Solution: 


D 


Exercise: 


Problem: 


What type of glial cell is the resident macrophage behind the blood- 
brain barrier? 


a. microglia 

b. astrocyte 

c. Schwann cell 
d. satellite cell 


Solution: 


A 
Exercise: 


Problem: 


What two types of macromolecules are the main components of 
myelin? 

a. carbohydrates and lipids 

b. proteins and nucleic acids 


c. lipids and proteins 
d. carbohydrates and nucleic acids 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: 


Multiple sclerosis is a demyelinating disease affecting the central 
nervous system. What type of cell would be the most likely target of 
this disease? Why? 


Solution: 

The disease would target oligodendrocytes. In the CNS, 

oligodendrocytes provide the myelin for axons. 
Exercise: 

Problem: 


Which type of neuron, based on its shape, is best suited for relaying 
information directly from one neuron to another? Explain why. 


Solution: 


Bipolar cells, because they have one dendrite that receives input and 
one axon that provides output, would be a direct relay between two 
other cells. 


Glossary 


astrocyte 
glial cell type of the CNS that provides support for neurons and 
maintains the blood-brain barrier 


axon hillock 
tapering of the neuron cell body that gives rise to the axon 


axon segment 
single stretch of the axon insulated by myelin and bounded by nodes of 
Ranvier at either end (except for the first, which is after the initial 
segment, and the last, which is followed by the axon terminal) 


axon terminal 
end of the axon, where there are usually several branches extending 
toward the target cell 


axoplasm 
cytoplasm of an axon, which is different in composition than the 
cytoplasm of the neuronal cell body 


bipolar 
shape of a neuron with two processes extending from the neuron cell 
body—the axon and one dendrite 


blood-brain barrier (BBB) 
physiological barrier between the circulatory system and the central 
nervous system that establishes a privileged blood supply, restricting 
the flow of substances into the CNS 


cerebrospinal fluid (CSF) 
circulatory medium within the CNS that is produced by ependymal 
cells in the choroid plexus filtering the blood 


choroid plexus 
specialized structure containing ependymal cells that line blood 
capillaries and filter blood to produce CSF in the four ventricles of the 
brain 


ependymal cell 
glial cell type in the CNS responsible for producing cerebrospinal fluid 


initial segment 
first part of the axon as it emerges from the axon hillock, where the 
electrical signals known as action potentials are generated 


microglia 
glial cell type in the CNS that serves as the resident component of the 


immune system 


multipolar 


shape of a neuron that has multiple processes—the axon and two or 
more dendrites 


myelin sheath 
lipid-rich layer of insulation that surrounds an axon, formed by 
oligodendrocytes in the CNS and Schwann cells in the PNS; facilitates 
the transmission of electrical signals 


node of Ranvier 
gap between two myelinated regions of an axon, allowing for 
strengthening of the electrical signal as it propagates down the axon 


oligodendrocyte 
glial cell type in the CNS that provides the myelin insulation for axons 
in tracts 


satellite cell 
glial cell type in the PNS that provides support for neurons in the 
ganglia 


Schwann cell 
glial cell type in the PNS that provides the myelin insulation for axons 
in nerves 


synapse 
narrow junction across which a chemical signal passes from neuron to 
the next, initiating a new electrical signal in the target cell 


synaptic end bulb 
swelling at the end of an axon where neurotransmitter molecules are 
released onto a target cell across a synapse 


unipolar 
shape of a neuron which has only one process that includes both the 
axon and dendrite 


ventricle 
central cavity within the brain where CSF is produced and circulates 


Anatomy of the CNS 
By the end of this section, you will be able to: 


e Name the major regions of the adult brain 

e Describe the connections between the cerebrum and brain stem 
through the diencephalon, and from those regions into the spinal cord 

e Recognize the complex connections within the subcortical structures 
of the basal nuclei 

e Explain the arrangement of gray and white matter in the spinal cord 


The brain and the spinal cord are the central nervous system, and they 
represent the main organs of the nervous system. The spinal cord is a single 
structure, whereas the adult brain is described in terms of four major 
regions: the cerebrum, the diencephalon, the brain stem, and the 
cerebellum. A person’s conscious experiences are based on neural activity 
in the brain. The regulation of homeostasis is governed by a specialized 
region in the brain. The coordination of reflexes depends on the integration 
of sensory and motor pathways in the spinal cord. 


The Cerebrum 


The iconic gray mantle of the human brain, which appears to make up most 
of the mass of the brain, is the cerebrum ((link]). The wrinkled portion is 
the cerebral cortex, and the rest of the structure is beneath that outer 
covering. There is a large separation between the two sides of the cerebrum 
called the longitudinal fissure. It separates the cerebrum into two distinct 
halves, a right and left cerebral hemisphere. Deep within the cerebrum, the 
white matter of the corpus callosum provides the major pathway for 
communication between the two hemispheres of the cerebral cortex. 

The Cerebrum 


Cerebrum 
Corpus callosum 


Longitudinal 
fissure 


Right hemisphere 


Cerebral cortex hemisphere 


Lateral view Anterior view 


The cerebrum is a large component of the CNS in humans, and the 
most obvious aspect of it is the folded surface called the cerebral 
cortex. 


Many of the higher neurological functions, such as memory, emotion, and 
consciousness, are the result of cerebral function. The complexity of the 
cerebrum is different across vertebrate species. The cerebrum of the most 
primitive vertebrates is not much more than the connection for the sense of 
smell. In mammals, the cerebrum comprises the outer gray matter that is the 
cortex (from the Latin word meaning “bark of a tree”) and several deep 
nuclei that belong to three important functional groups. The basal nuclei 
are responsible for cognitive processing, the most important function being 
that associated with planning movements. The basal forebrain contains 
nuclei that are important in learning and memory. The limbic cortex is the 
region of the cerebral cortex that is part of the limbic system, a collection 
of structures involved in emotion, memory, and behavior. 


Cerebral Cortex 


The cerebrum is covered by a continuous layer of gray matter that wraps 
around either side of the forebrain—the cerebral cortex. This thin, extensive 
region of wrinkled gray matter is responsible for the higher functions of the 
nervous system. A gyrus (plural = gyri) is the ridge of one of those 
wrinkles, and a sulcus (plural = sulci) is the groove between two gyri. The 
pattern of these folds of tissue indicates specific regions of the cerebral 
cortex. 


The head is limited by the size of the birth canal, and the brain must fit 
inside the cranial cavity of the skull. Extensive folding in the cerebral 
cortex enables more gray matter to fit into this limited space. If the gray 
matter of the cortex were peeled off of the cerebrum and laid out flat, its 
surface area would be roughly equal to one square meter. 


The folding of the cortex maximizes the amount of gray matter in the 
cranial cavity. During embryonic development, as the telencephalon 
expands within the skull, the brain goes through a regular course of growth 
that results in everyone’s brain having a similar pattern of folds. The surface 
of the brain can be mapped on the basis of the locations of large gyri and 
sulci. Using these landmarks, the cortex can be separated into four major 
regions, or lobes ((link]). The lateral sulcus that separates the temporal 
lobe from the other regions is one such landmark. Superior to the lateral 
sulcus are the parietal lobe and frontal lobe, which are separated from 
each other by the central sulcus. The posterior region of the cortex is the 
occipital lobe, which has no obvious anatomical border between it and the 
parietal or temporal lobes on the lateral surface of the brain. From the 
medial surface, an obvious landmark separating the parietal and occipital 
lobes is called the parieto-occipital sulcus. The fact that there is no 
obvious anatomical border between these lobes is consistent with the 
functions of these regions being interrelated. 

Lobes of the Cerebral Cortex 


Central sulcus 


Precentral gyrus Postcentral gyrus 


Frontal lobe Parietal lobe 


Parieto-occipital 
sulcus 


Lateral sulcus 


Occipital lobe 


Temporal lobe 


The cerebral cortex is divided into four 
lobes. Extensive folding increases the 
surface area available for cerebral 
functions. 


Different regions of the cerebral cortex can be associated with particular 
functions, a concept known as localization of function. In the early 1900s, a 
German neuroscientist named Korbinian Brodmann performed an extensive 
study of the microscopic anatomy—the cytoarchitecture—of the cerebral 
cortex and divided the cortex into 52 separate regions on the basis of the 
histology of the cortex. His work resulted in a system of classification 
known as Brodmann’s areas, which is still used today to describe the 
anatomical distinctions within the cortex ({link]). The results from 
Brodmann’s work on the anatomy align very well with the functional 
differences within the cortex. Areas 17 and 18 in the occipital lobe are 
responsible for primary visual perception. That visual information is 
complex, so it is processed in the temporal and parietal lobes as well. 


The temporal lobe is associated with primary auditory sensation, known as 
Brodmann’s areas 41 and 42 in the superior temporal lobe. Because regions 


of the temporal lobe are part of the limbic system, memory is an important 
function associated with that lobe. Memory is essentially a sensory 
function; memories are recalled sensations such as the smell of Mom’s 
baking or the sound of a barking dog. Even memories of movement are 
really the memory of sensory feedback from those movements, such as 
stretching muscles or the movement of the skin around a joint. Structures in 
the temporal lobe are responsible for establishing long-term memory, but 
the ultimate location of those memories is usually in the region in which the 
sensory perception was processed. 


The main sensation associated with the parietal lobe is somatosensation, 
meaning the general sensations associated with the body. Posterior to the 
central sulcus is the postcentral gyrus, the primary somatosensory cortex, 
which is identified as Brodmann’s areas 1, 2, and 3. All of the tactile senses 
are processed in this area, including touch, pressure, tickle, pain, itch, and 
vibration, as well as more general senses of the body such as 
proprioception and kinesthesia, which are the senses of body position and 
movement, respectively. 


Anterior to the central sulcus is the frontal lobe, which is primarily 
associated with motor functions. The precentral gyrus is the primary 
motor cortex. Cells from this region of the cerebral cortex are the upper 
motor neurons that instruct cells in the spinal cord to move skeletal 
muscles. Anterior to this region are a few areas that are associated with 
planned movements. The premotor area is responsible for thinking of a 
movement to be made. The frontal eye fields are important in eliciting eye 
movements and in attending to visual stimuli. Broca’s area is responsible 
for the production of language, or controlling movements responsible for 
speech; in the vast majority of people, it is located only on the left side. 
Anterior to these regions is the prefrontal lobe, which serves cognitive 
functions that can be the basis of personality, short-term memory, and 
consciousness. The prefrontal lobotomy is an outdated mode of treatment 
for personality disorders (psychiatric conditions) that profoundly affected 
the personality of the patient. 

Brodmann's Areas of the Cerebral Cortex 


Areas 1, 2,3 
Primary 

somatosensory 
cortex 


Area 4 
Primary motor cortex 


Areas 44, 45 
Broca’s area 


Area 4 
Primary motor 
cortex 


Areas 39, 40 
Wernicke’s area 


d 


Area 22 % Area 17 


: ; SB oe ee ok A : 
Primary auditory “Sel fe 2» Primary visual cortex 
cortex 


Brodmann’s cytotechtonic map (1909): Brodmann’s cytotechtonic map (1909): 
Lateral surface Medial surface 


Brodmann mapping of functionally distinct regions of the 
cortex was based on its cytoarchitecture at a microscopic level. 


Subcortical structures 


Beneath the cerebral cortex are sets of nuclei known as subcortical nuclei 
that augment cortical processes. The nuclei of the basal forebrain serve as 
the primary location for acetylcholine production, which modulates the 
overall activity of the cortex, possibly leading to greater attention to sensory 
stimuli. Alzheimer’s disease is associated with a loss of neurons in the basal 
forebrain. The hippocampus and amygdala are medial-lobe structures that, 
along with the adjacent cortex, are involved in long-term memory formation 
and emotional responses. The basal nuclei are a set of nuclei in the 
cerebrum responsible for comparing cortical processing with the general 
state of activity in the nervous system to influence the likelihood of 
movement taking place. For example, while a student is sitting in a 
classroom listening to a lecture, the basal nuclei will keep the urge to jump 
up and scream from actually happening. (The basal nuclei are also referred 


to as the basal ganglia, although that is potentially confusing because the 
term ganglia is typically used for peripheral structures.) 


The major structures of the basal nuclei that control movement are the 
caudate, putamen, and globus pallidus, which are located deep in the 
cerebrum. The caudate is a long nucleus that follows the basic C-shape of 
the cerebrum from the frontal lobe, through the parietal and occipital lobes, 
into the temporal lobe. The putamen is mostly deep in the anterior regions 
of the frontal and parietal lobes. Together, the caudate and putamen are 
called the striatum. The globus pallidus is a layered nucleus that lies just 
medial to the putamen; they are called the lenticular nuclei because they 
look like curved pieces fitting together like lenses. The globus pallidus has 
two subdivisions, the external and internal segments, which are lateral and 
medial, respectively. These nuclei are depicted in a frontal section of the 
brain in [link]. 

Frontal Section of Cerebral Cortex and Basal Nuclei 


Lateral ventricle 


Striatum: 
Caudate 
Putamen 

Corpus 

callosum 


Globus pallidus 
Frontal section 


The major components of the basal 
nuclei, shown in a frontal section of the 
brain, are the caudate (just lateral to the 

lateral ventricle), the putamen (inferior to 
the caudate and separated by the large 
white-matter structure called the internal 


capsule), and the globus pallidus (medial 
to the putamen). 


The basal nuclei in the cerebrum are connected with a few more nuclei in 
the brain stem that together act as a functional group that forms a motor 
pathway. Two streams of information processing take place in the basal 
nuclei. All input to the basal nuclei is from the cortex into the striatum 
({link]). The direct pathway is the projection of axons from the striatum to 
the globus pallidus internal segment (GPi) and the substantia nigra pars 
reticulata (SNr). The GPi/SNr then projects to the thalamus, which projects 
back to the cortex. The indirect pathway is the projection of axons from 
the striatum to the globus pallidus external segment (GPe), then to the 
subthalamic nucleus (STN), and finally to GPi/SNr. The two streams both 
target the GPi/SNr, but one has a direct projection and the other goes 
through a few intervening nuclei. The direct pathway causes the 
disinhibition of the thalamus (inhibition of one cell on a target cell that 
then inhibits the first cell), whereas the indirect pathway causes, or 
reinforces, the normal inhibition of the thalamus. The thalamus then can 
either excite the cortex (as a result of the direct pathway) or fail to excite 
the cortex (as a result of the indirect pathway). 


Connections of Basal Nuclei 
Basal nuclei 


} t = Glutamate 
= Dopamine 


A 


i 


[Thalarws |<—] 


Input to the basal nuclei is from 
the cerebral cortex, which is an 


excitatory connection releasing 
glutamate as a 
neurotransmitter. This input is 
to the striatum, or the caudate 
and putamen. In the direct 
pathway, the striatum projects 
to the internal segment of the 
globus pallidus and the 
substantia nigra pars reticulata 
(GPi/SNr). This is an inhibitory 
pathway, in which GABA is 
released at the synapse, and the 
target cells are hyperpolarized 
and less likely to fire. The 
output from the basal nuclei is 
to the thalamus, which is an 
inhibitory projection using 
GABA. 


The switch between the two pathways is the substantia nigra pars 
compacta, which projects to the striatum and releases the neurotransmitter 
dopamine. Dopamine receptors are either excitatory (D1-type receptors) or 
inhibitory (D2-type receptors). The direct pathway is activated by 
dopamine, and the indirect pathway is inhibited by dopamine. When the 
substantia nigra pars compacta is firing, it signals to the basal nuclei that the 
body is in an active state, and movement will be more likely. When the 
substantia nigra pars compacta is silent, the body is in a passive state, and 
movement is inhibited. To illustrate this situation, while a student is sitting 
listening to a lecture, the substantia nigra pars compacta would be silent and 
the student less likely to get up and walk around. Likewise, while the 
professor is lecturing, and walking around at the front of the classroom, the 
professor’s substantia nigra pars compacta would be active, in keeping with 
his or her activity level. 


Watch this video to learn about the basal nuclei (also known as the basal 
ganglia), which have two pathways that process information within the 
cerebrum. As shown in this video, the direct pathway is the shorter 
pathway through the system that results in increased activity in the cerebral 
cortex and increased motor activity. The direct pathway is described as 
resulting in “disinhibition” of the thalamus. What does disinhibition mean? 
What are the two neurons doing individually to cause this? 


Note: 


[elie 


— 
mss Openstax COLLEGE 
ro-e." 


Watch this video to learn about the basal nuclei (also known as the basal 
ganglia), which have two pathways that process information within the 
cerebrum. As shown in this video, the indirect pathway is the longer 
pathway through the system that results in decreased activity in the 
cerebral cortex, and therefore less motor activity. The indirect pathway has 
an extra couple of connections in it, including disinhibition of the 
subthalamic nucleus. What is the end result on the thalamus, and therefore 
on movement initiated by the cerebral cortex? 


Note: 

Everyday Connections 

The Myth of Left Brain/Right Brain 

There is a persistent myth that people are “right-brained” or “left-brained,” 
which is an oversimplification of an important concept about the cerebral 
hemispheres. There is some lateralization of function, in which the left side 
of the brain is devoted to language function and the right side is devoted to 
spatial and nonverbal reasoning. Whereas these functions are 
predominantly associated with those sides of the brain, there is no 
monopoly by either side on these functions. Many pervasive functions, 
such as language, are distributed globally around the cerebrum. 

Some of the support for this misconception has come from studies of split 
brains. A drastic way to deal with a rare and devastating neurological 
condition (intractable epilepsy) is to separate the two hemispheres of the 
brain. After sectioning the corpus callosum, a split-brained patient will 
have trouble producing verbal responses on the basis of sensory 
information processed on the right side of the cerebrum, leading to the idea 
that the left side is responsible for language function. 

However, there are well-documented cases of language functions lost from 
damage to the right side of the brain. The deficits seen in damage to the left 
side of the brain are classified as aphasia, a loss of speech function; 
damage on the right side can affect the use of language. Right-side damage 
can result in a loss of ability to understand figurative aspects of speech, 
such as jokes, irony, or metaphors. Nonverbal aspects of speech can be 
affected by damage to the right side, such as facial expression or body 
language, and right-side damage can lead to a “flat affect” in speech, or a 
loss of emotional expression in speech—sounding like a robot when 
talking. 


The Diencephalon 


The diencephalon is the one region of the adult brain that retains its name 
from embryologic development. The etymology of the word diencephalon 
translates to “through brain.” It is the connection between the cerebrum and 
the rest of the nervous system, with one exception. The rest of the brain, the 


spinal cord, and the PNS all send information to the cerebrum through the 
diencephalon. Output from the cerebrum passes through the diencephalon. 
The single exception is the system associated with olfaction, or the sense of 
smell, which connects directly with the cerebrum. In the earliest vertebrate 
species, the cerebrum was not much more than olfactory bulbs that received 
peripheral information about the chemical environment (to call it smell in 
these organisms is imprecise because they lived in the ocean). 


The diencephalon is deep beneath the cerebrum and constitutes the walls of 
the third ventricle. The diencephalon can be described as any region of the 
brain with “thalamus” in its name. The two major regions of the 
diencephalon are the thalamus itself and the hypothalamus ([link]). There 
are other structures, such as the epithalamus, which contains the pineal 
gland, or the subthalamus, which includes the subthalamic nucleus that is 
part of the basal nuclei. 


Thalamus 


The thalamus is a collection of nuclei that relay information between the 
cerebral cortex and the periphery, spinal cord, or brain stem. All sensory 
information, except for the sense of smell, passes through the thalamus 
before processing by the cortex. Axons from the peripheral sensory organs, 
or intermediate nuclei, synapse in the thalamus, and thalamic neurons 
project directly to the cerebrum. It is a requisite synapse in any sensory 
pathway, except for olfaction. The thalamus does not just pass the 
information on, it also processes that information. For example, the portion 
of the thalamus that receives visual information will influence what visual 
stimuli are important, or what receives attention. 


The cerebrum also sends information down to the thalamus, which usually 
communicates motor commands. This involves interactions with the 
cerebellum and other nuclei in the brain stem. The cerebrum interacts with 
the basal nuclei, which involves connections with the thalamus. The 
primary output of the basal nuclei is to the thalamus, which relays that 
output to the cerebral cortex. The cortex also sends information to the 
thalamus that will then influence the effects of the basal nuclei. 


Hypothalamus 


Inferior and slightly anterior to the thalamus is the hypothalamus, the other 
major region of the diencephalon. The hypothalamus is a collection of 
nuclei that are largely involved in regulating homeostasis. The 
hypothalamus is the executive region in charge of the autonomic nervous 
system and the endocrine system through its regulation of the anterior 
pituitary gland. Other parts of the hypothalamus are involved in memory 
and emotion as part of the limbic system. 

The Diencephalon 


Thalamus 


Hypothalamus 


Pituitary gland 


The diencephalon is composed primarily of the 
thalamus and hypothalamus, which together define 
the walls of the third ventricle. The thalami are 
two elongated, ovoid structures on either side of 
the midline that make contact in the middle. The 
hypothalamus is inferior and anterior to the 
thalamus, culminating in a sharp angle to which 
the pituitary gland is attached. 


Brain Stem 


The midbrain and hindbrain (composed of the pons and the medulla) are 
collectively referred to as the brain stem ({link]). The structure emerges 
from the ventral surface of the forebrain as a tapering cone that connects the 
brain to the spinal cord. Attached to the brain stem, but considered a 
separate region of the adult brain, is the cerebellum. The midbrain 
coordinates sensory representations of the visual, auditory, and 
somatosensory perceptual spaces. The pons is the main connection with the 
cerebellum. The pons and the medulla regulate several crucial functions, 
including the cardiovascular and respiratory systems and rates. 


The cranial nerves connect through the brain stem and provide the brain 
with the sensory input and motor output associated with the head and neck, 
including most of the special senses. The major ascending and descending 
pathways between the spinal cord and brain, specifically the cerebrum, pass 
through the brain stem. 

The Brain Stem 


Midbrain 


Pons 


Medulla 


The brain stem comprises three regions: the 
midbrain, the pons, and the medulla. 


Midbrain 


One of the original regions of the embryonic brain, the midbrain is a small 
region between the thalamus and pons. It is separated into the tectum and 
tegmentum, from the Latin words for roof and floor, respectively. The 
cerebral aqueduct passes through the center of the midbrain, such that these 
regions are the roof and floor of that canal. 


The tectum is composed of four bumps known as the colliculi (singular = 
colliculus), which means “little hill” in Latin. The inferior colliculus is the 
inferior pair of these enlargements and is part of the auditory brain stem 
pathway. Neurons of the inferior colliculus project to the thalamus, which 
then sends auditory information to the cerebrum for the conscious 
perception of sound. The superior colliculus is the superior pair and 
combines sensory information about visual space, auditory space, and 
somatosensory space. Activity in the superior colliculus is related to 
orienting the eyes to a sound or touch stimulus. If you are walking along the 
sidewalk on campus and you hear chirping, the superior colliculus 
coordinates that information with your awareness of the visual location of 
the tree right above you. That is the correlation of auditory and visual maps. 
If you suddenly feel something wet fall on your head, your superior 
colliculus integrates that with the auditory and visual maps and you know 
that the chirping bird just relieved itself on you. You want to look up to see 
the culprit, but do not. 


The tegmentum is continuous with the gray matter of the rest of the brain 
stem. Throughout the midbrain, pons, and medulla, the tegmentum contains 
the nuclei that receive and send information through the cranial nerves, as 
well as regions that regulate important functions such as those of the 
cardiovascular and respiratory systems. 


Pons 


The word pons comes from the Latin word for bridge. It is visible on the 
anterior surface of the brain stem as the thick bundle of white matter 
attached to the cerebellum. The pons is the main connection between the 
cerebellum and the brain stem. The bridge-like white matter is only the 
anterior surface of the pons; the gray matter beneath that is a continuation 
of the tegmentum from the midbrain. Gray matter in the tegmentum region 
of the pons contains neurons receiving descending input from the forebrain 
that is sent to the cerebellum. 


Medulla 


The medulla is the region known as the myelencephalon in the embryonic 
brain. The initial portion of the name, “myel,” refers to the significant white 
matter found in this region—especially on its exterior, which is continuous 
with the white matter of the spinal cord. The tegmentum of the midbrain 
and pons continues into the medulla because this gray matter is responsible 
for processing cranial nerve information. A diffuse region of gray matter 
throughout the brain stem, known as the reticular formation, is related to 
sleep and wakefulness, such as general brain activity and attention. 


The Cerebellum 


The cerebellum, as the name suggests, is the “little brain.” It is covered in 
gyri and sulci like the cerebrum, and looks like a miniature version of that 
part of the brain ([link]). The cerebellum is largely responsible for 
comparing information from the cerebrum with sensory feedback from the 
periphery through the spinal cord. It accounts for approximately 10 percent 
of the mass of the brain. 

The Cerebellum 


Cerebellum 


Deep cerebellar 
white matter 
(arbor vitae) 


Inferior olive 


The cerebellum is situated on the posterior 
surface of the brain stem. Descending input 
from the cerebellum enters through the large 

white matter structure of the pons. Ascending 

input from the periphery and spinal cord enters 

through the fibers of the inferior olive. Output 
goes to the midbrain, which sends a 
descending signal to the spinal cord. 


Descending fibers from the cerebrum have branches that connect to neurons 
in the pons. Those neurons project into the cerebellum, providing a copy of 


motor commands sent to the spinal cord. Sensory information from the 
periphery, which enters through spinal or cranial nerves, is copied to a 
nucleus in the medulla known as the inferior olive. Fibers from this nucleus 
enter the cerebellum and are compared with the descending commands 
from the cerebrum. If the primary motor cortex of the frontal lobe sends a 
command down to the spinal cord to initiate walking, a copy of that 
instruction is sent to the cerebellum. Sensory feedback from the muscles 
and joints, proprioceptive information about the movements of walking, and 
sensations of balance are sent to the cerebellum through the inferior olive 
and the cerebellum compares them. If walking is not coordinated, perhaps 
because the ground is uneven or a strong wind is blowing, then the 
cerebellum sends out a corrective command to compensate for the 
difference between the original cortical command and the sensory feedback. 
The output of the cerebellum is into the midbrain, which then sends a 
descending input to the spinal cord to correct the messages going to skeletal 
muscles. 


The Spinal Cord 


The description of the CNS is concentrated on the structures of the brain, 
but the spinal cord is another major organ of the system. Whereas the brain 
develops out of expansions of the neural tube into primary and then 
secondary vesicles, the spinal cord maintains the tube structure and is only 
specialized into certain regions. As the spinal cord continues to develop in 
the newborn, anatomical features mark its surface. The anterior midline is 
marked by the anterior median fissure, and the posterior midline is 
marked by the posterior median sulcus. Axons enter the posterior side 
through the dorsal (posterior) nerve root, which marks the posterolateral 
sulcus on either side. The axons emerging from the anterior side do so 
through the ventral (anterior) nerve root. Note that it is common to see 
the terms dorsal (dorsal = “back”) and ventral (ventral = “belly”) used 
interchangeably with posterior and anterior, particularly in reference to 
nerves and the structures of the spinal cord. You should learn to be 
comfortable with both. 


On the whole, the posterior regions are responsible for sensory functions 
and the anterior regions are associated with motor functions. This comes 


from the initial development of the spinal cord, which is divided into the 
basal plate and the alar plate. The basal plate is closest to the ventral 
midline of the neural tube, which will become the anterior face of the spinal 
cord and gives rise to motor neurons. The alar plate is on the dorsal side of 
the neural tube and gives rise to neurons that will receive sensory input 
from the periphery. 


The length of the spinal cord is divided into regions that correspond to the 
regions of the vertebral column. The name of a spinal cord region 
corresponds to the level at which spinal nerves pass through the 
intervertebral foramina. Immediately adjacent to the brain stem is the 
cervical region, followed by the thoracic, then the lumbar, and finally the 
sacral region. The spinal cord is not the full length of the vertebral column 
because the spinal cord does not grow significantly longer after the first or 
second year, but the skeleton continues to grow. The nerves that emerge 
from the spinal cord pass through the intervertebral formina at the 
respective levels. As the vertebral column grows, these nerves grow with it 
and result in a long bundle of nerves that resembles a horse’s tail and is 
named the cauda equina. The sacral spinal cord is at the level of the upper 
lumbar vertebral bones. The spinal nerves extend from their various levels 
to the proper level of the vertebral column. 


Gray Horns 


In cross-section, the gray matter of the spinal cord has the appearance of an 
ink-blot test, with the spread of the gray matter on one side replicated on the 
other—a shape reminiscent of a bulbous capital “H.” As shown in [link], 
the gray matter is subdivided into regions that are referred to as horns. The 
posterior horn is responsible for sensory processing. The anterior horn 
sends out motor signals to the skeletal muscles. The lateral horn, which is 
only found in the thoracic, upper lumbar, and sacral regions, is the central 
component of the sympathetic division of the autonomic nervous system. 


Some of the largest neurons of the spinal cord are the multipolar motor 
neurons in the anterior horn. The fibers that cause contraction of skeletal 
muscles are the axons of these neurons. The motor neuron that causes 


contraction of the big toe, for example, is located in the sacral spinal cord. 
The axon that has to reach all the way to the belly of that muscle may be a 
meter in length. The neuronal cell body that maintains that long fiber must 
be quite large, possibly several hundred micrometers in diameter, making it 
one of the largest cells in the body. 

Cross-section of Spinal Cord 


Posterior (dorsal) 
columns 


Gray matter: 
Posterior (dorsal) ~¢ 
horn 


Lateral columns 


Lateral horn 
Central canal 


Anterior (ventral) 


Anterior (ventral) columns 
horn 


The cross-section of a thoracic spinal cord 
segment shows the posterior, anterior, and 
lateral horns of gray matter, as well as the 
posterior, anterior, and lateral columns of 
white matter. LM x 40. (Micrograph 
provided by the Regents of University of 
Michigan Medical School © 2012) 


White Columns 


Just as the gray matter is separated into horns, the white matter of the spinal 
cord is separated into columns. Ascending tracts of nervous system fibers 
in these columns carry sensory information up to the brain, whereas 
descending tracts carry motor commands from the brain. Looking at the 
spinal cord longitudinally, the columns extend along its length as 
continuous bands of white matter. Between the two posterior horns of gray 
matter are the posterior columns. Between the two anterior horns, and 
bounded by the axons of motor neurons emerging from that gray matter 
area, are the anterior columns. The white matter on either side of the 
spinal cord, between the posterior horn and the axons of the anterior horn 
neurons, are the lateral columns. The posterior columns are composed of 
axons of ascending tracts. The anterior and lateral columns are composed of 
many different groups of axons of both ascending and descending tracts— 
the latter carrying motor commands down from the brain to the spinal cord 
to control output to the periphery. 


Note: 


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Watch this video to learn about the gray matter of the spinal cord that 
receives input from fibers of the dorsal (posterior) root and sends 
information out through the fibers of the ventral (anterior) root. As 
discussed in this video, these connections represent the interactions of the 
CNS with peripheral structures for both sensory and motor functions. The 
cervical and lumbar spinal cords have enlargements as a result of larger 
populations of neurons. What are these enlargements responsible for? 


Note: 

Disorders of the... 

Basal Nuclei 

Parkinson’s disease is a disorder of the basal nuclei, specifically of the 
substantia nigra, that demonstrates the effects of the direct and indirect 
pathways. Parkinson’s disease is the result of neurons in the substantia 
nigra pars compacta dying. These neurons release dopamine into the 
striatum. Without that modulatory influence, the basal nuclei are stuck in 
the indirect pathway, without the direct pathway being activated. The direct 
pathway is responsible for increasing cortical movement commands. The 
increased activity of the indirect pathway results in the hypokinetic 
disorder of Parkinson’s disease. 

Parkinson’s disease is neurodegenerative, meaning that neurons die that 
cannot be replaced, so there is no cure for the disorder. Treatments for 
Parkinson’s disease are aimed at increasing dopamine levels in the 
striatum. Currently, the most common way of doing that is by providing 
the amino acid L-DOPA, which is a precursor to the neurotransmitter 
dopamine and can cross the blood-brain barrier. With levels of the 
precursor elevated, the remaining cells of the substantia nigra pars 
compacta can make more neurotransmitter and have a greater effect. 
Unfortunately, the patient will become less responsive to L-DOPA 
treatment as time progresses, and it can cause increased dopamine levels 
elsewhere in the brain, which are associated with psychosis or 
schizophrenia. 


Note: 
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=> Openstax COLLEGE 
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Visit this site for a thorough explanation of Parkinson’s disease. 


Note: 


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meee OPENStAX COLLEGE 


Compared with the nearest evolutionary relative, the chimpanzee, the 
human has a brain that is huge. At a point in the past, a common ancestor 
gave rise to the two species of humans and chimpanzees. That evolutionary 
history is long and is still an area of intense study. But something happened 
to increase the size of the human brain relative to the chimpanzee. Read 
this article in which the author explores the current understanding of why 
this happened. 

According to one hypothesis about the expansion of brain size, what tissue 
might have been sacrificed so energy was available to grow our larger 
brain? Based on what you know about that tissue and nervous tissue, why 
would there be a trade-off between them in terms of energy use? 


Chapter Review 


The adult brain is separated into four major regions: the cerebrum, the 
diencephalon, the brain stem, and the cerebellum. The cerebrum is the 
largest portion and contains the cerebral cortex and subcortical nuclei. It is 
divided into two halves by the longitudinal fissure. 


The cortex is separated into the frontal, parietal, temporal, and occipital 
lobes. The frontal lobe is responsible for motor functions, from planning 
movements through executing commands to be sent to the spinal cord and 
periphery. The most anterior portion of the frontal lobe is the prefrontal 
cortex, which is associated with aspects of personality through its influence 
on motor responses in decision-making. 


The other lobes are responsible for sensory functions. The parietal lobe is 
where somatosensation is processed. The occipital lobe is where visual 
processing begins, although the other parts of the brain can contribute to 
visual function. The temporal lobe contains the cortical area for auditory 
processing, but also has regions crucial for memory formation. 


Nuclei beneath the cerebral cortex, known as the subcortical nuclei, are 
responsible for augmenting cortical functions. The basal nuclei receive 
input from cortical areas and compare it with the general state of the 
individual through the activity of a dopamine-releasing nucleus. The output 
influences the activity of part of the thalamus that can then increase or 
decrease cortical activity that often results in changes to motor commands. 
The basal forebrain is responsible for modulating cortical activity in 
attention and memory. The limbic system includes deep cerebral nuclei that 
are responsible for emotion and memory. 


The diencephalon includes the thalamus and the hypothalamus, along with 
some other structures. The thalamus is a relay between the cerebrum and 
the rest of the nervous system. The hypothalamus coordinates homeostatic 
functions through the autonomic and endocrine systems. 


The brain stem is composed of the midbrain, pons, and medulla. It controls 
the head and neck region of the body through the cranial nerves. There are 
control centers in the brain stem that regulate the cardiovascular and 
respiratory systems. 


The cerebellum is connected to the brain stem, primarily at the pons, where 
it receives a copy of the descending input from the cerebrum to the spinal 
cord. It can compare this with sensory feedback input through the medulla 
and send output through the midbrain that can correct motor commands for 
coordination. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to learn about the basal nuclei (also known as the 
basal ganglia), which have two pathways that process information 
within the cerebrum. As shown in this video, the direct pathway is the 
shorter pathway through the system that results in increased activity in 
the cerebral cortex and increased motor activity. The direct pathway is 
described as resulting in “disinhibition” of the thalamus. What does 
disinhibition mean? What are the two neurons doing individually to 
cause this? 


Solution: 


Both cells are inhibitory. The first cell inhibits the second one. 
Therefore, the second cell can no longer inhibit its target. This is 
disinhibition of that target across two synapses. 


Exercise: 


Problem: 


Watch this video to learn about the basal nuclei (also known as the 
basal ganglia), which have two pathways that process information 
within the cerebrum. As shown in this video, the indirect pathway is 
the longer pathway through the system that results in decreased 
activity in the cerebral cortex, and therefore less motor activity. The 
indirect pathway has an extra couple of connections in it, including 
disinhibition of the subthalamic nucleus. What is the end result on the 
thalamus, and therefore on movement initiated by the cerebral cortex? 


Solution: 


By disinhibiting the subthalamic nucleus, the indirect pathway 
increases excitation of the globus pallidus internal segment. That, in 
turn, inhibits the thalamus, which is the opposite effect of the direct 
pathway that disinhibits the thalamus. 


Exercise: 


Problem: 


Watch this video to learn about the gray matter of the spinal cord that 
receives input from fibers of the dorsal (posterior) root and sends 
information out through the fibers of the ventral (anterior) root. As 
discussed in this video, these connections represent the interactions of 
the CNS with peripheral structures for both sensory and motor 
functions. The cervical and lumbar spinal cords have enlargements as a 
result of larger populations of neurons. What are these enlargements 
responsible for? 


Solution: 


There are more motor neurons in the anterior horns that are responsible 
for movement in the limbs. The cervical enlargement is for the arms, 
and the lumbar enlargement is for the legs. 


Exercise: 


Problem: 


Compared with the nearest evolutionary relative, the chimpanzee, the 
human has a brain that is huge. At a point in the past, a common 
ancestor gave rise to the two species of humans and chimpanzees. That 
evolutionary history is long and is still an area of intense study. But 
something happened to increase the size of the human brain relative to 
the chimpanzee. Read this article in which the author explores the 
current understanding of why this happened. 


According to one hypothesis about the expansion of brain size, what 
tissue might have been sacrificed so energy was available to grow our 
larger brain? Based on what you know about that tissue and nervous 
tissue, why would there be a trade-off between them in terms of energy 
use? 


Solution: 


Energy is needed for the brain to develop and perform higher cognitive 
functions. That energy is not available for the muscle tissues to 
develop and function. The hypothesis suggests that humans have larger 
brains and less muscle mass, and chimpanzees have the smaller brains 
but more muscle mass. 


Review Questions 
Exercise: 


Problem: 


Which lobe of the cerebral cortex is responsible for generating motor 
commands? 


a. temporal 
b. parietal 
c. occipital 
d. frontal 


Solution: 
D 
Exercise: 
Problem: What region of the diencephalon coordinates homeostasis? 
a. thalamus 
b. epithalamus 


c. hypothalamus 
d. subthalamus 


Solution: 


C 


Exercise: 


Problem: 
What level of the brain stem is the major input to the cerebellum? 


a. midbrain 
b. pons 

c. medulla 

d. spinal cord 


Solution: 


B 
Exercise: 


Problem: 


What region of the spinal cord contains motor neurons that direct the 
movement of skeletal muscles? 


a. anterior horn 
b. posterior horn 
c. lateral horn 
d. alar plate 


Solution: 


A 
Exercise: 


Problem: 


Brodmann’s areas map different regions of the to particular 
functions. 


a. cerebellum 


b. cerebral cortex 
c. basal forebrain 
d. corpus callosum 


Solution: 


B 


Critical Thinking Questions 


Exercise: 
Problem: 
Damage to specific regions of the cerebral cortex, such as through a 


stroke, can result in specific losses of function. What functions would 
likely be lost by a stroke in the temporal lobe? 


Solution: 


The temporal lobe has sensory functions associated with hearing and 
vision, as well as being important for memory. A stroke in the 
temporal lobe can result in specific sensory deficits in these systems 
(known as agnosias) or losses in memory. 


Exercise: 
Problem: 


Why do the anatomical inputs to the cerebellum suggest that it can 
compare motor commands and sensory feedback? 


Solution: 


A copy of descending input from the cerebrum to the spinal cord, 
through the pons, and sensory feedback from the spinal cord and 
special senses like balance, through the medulla, both go to the 


cerebellum. It can therefore send output through the midbrain that will 
correct spinal cord control of skeletal muscle movements. 


Glossary 


alar plate 
developmental region of the spinal cord that gives rise to the posterior 
horn of the gray matter 


amygdala 
nucleus deep in the temporal lobe of the cerebrum that is related to 
memory and emotional behavior 


anterior column 
white matter between the anterior horns of the spinal cord composed of 
many different groups of axons of both ascending and descending 
tracts 


anterior horn 
gray matter of the spinal cord containing multipolar motor neurons, 
sometimes referred to as the ventral horn 


anterior median fissure 
deep midline feature of the anterior spinal cord, marking the separation 
between the right and left sides of the cord 


ascending tract 
central nervous system fibers carrying sensory information from the 
spinal cord or periphery to the brain 


basal forebrain 
nuclei of the cerebrum related to modulation of sensory stimuli and 
attention through broad projections to the cerebral cortex, loss of 
which is related to Alzheimer’s disease 


basal nuclei 


nuclei of the cerebrum (with a few components in the upper brain stem 
and diencephalon) that are responsible for assessing cortical movement 
commands and comparing them with the general state of the individual 
through broad modulatory activity of dopamine neurons; largely 
related to motor functions, as evidenced through the symptoms of 
Parkinson’s and Huntington’s diseases 


basal plate 
developmental region of the spinal cord that gives rise to the lateral 
and anterior horns of gray matter 


Broca’s area 
region of the frontal lobe associated with the motor commands 
necessary for speech production and located only in the cerebral 
hemisphere responsible for language production, which is the left side 
in approximately 95 percent of the population 


Brodmann’s areas 
mapping of regions of the cerebral cortex based on microscopic 
anatomy that relates specific areas to functional differences, as 
described by Brodmann in the early 1900s 


cauda equina 
bundle of spinal nerve roots that descend from the lower spinal cord 
below the first lumbar vertebra and lie within the vertebral cavity; has 
the appearance of a horse's tail 


caudate 
nucleus deep in the cerebrum that is part of the basal nuclei; along with 
the putamen, it is part of the striatum 


central sulcus 
surface landmark of the cerebral cortex that marks the boundary 
between the frontal and parietal lobes 


cerebral cortex 
outer gray matter covering the forebrain, marked by wrinkles and folds 
known as gyri and sulci 


cerebrum 
region of the adult brain that develops from the telencephalon and is 
responsible for higher neurological functions such as memory, 
emotion, and consciousness 


cerebellum 
region of the adult brain connected primarily to the pons that 
developed from the metencephalon (along with the pons) and is largely 
responsible for comparing information from the cerebrum with sensory 
feedback from the periphery through the spinal cord 


cerebral hemisphere 
one half of the bilaterally symmetrical cerebrum 


corpus callosum 
large white matter structure that connects the right and left cerebral 
hemispheres 


descending tract 
central nervous system fibers carrying motor commands from the brain 
to the spinal cord or periphery 


direct pathway 
connections within the basal nuclei from the striatum to the globus 
pallidus internal segment and substantia nigra pars reticulata that 
disinhibit the thalamus to increase cortical control of movement 


disinhibition 
disynaptic connection in which the first synapse inhibits the second 


cell, which then stops inhibiting the final target 


dorsal (posterior) nerve root 
axons entering the posterior horn of the spinal cord 


epithalamus 
region of the diecephalon containing the pineal gland 


frontal eye field 


region of the frontal lobe associated with motor commands to orient 
the eyes toward an object of visual attention 


frontal lobe 
region of the cerebral cortex directly beneath the frontal bone of the 
cranium 


globus pallidus 
nuclei deep in the cerebrum that are part of the basal nuclei and can be 
divided into the internal and external segments 


gyrus 
ridge formed by convolutions on the surface of the cerebrum or 


cerebellum 


hippocampus 
gray matter deep in the temporal lobe that is very important for long- 
term memory formation 


hypothalamus 
major region of the diencephalon that is responsible for coordinating 
autonomic and endocrine control of homeostasis 


indirect pathway 
connections within the basal nuclei from the striatum through the 
globus pallidus external segment and subthalamic nucleus to the 
globus pallidus internal segment/substantia nigra pars compacta that 
result in inhibition of the thalamus to decrease cortical control of 
movement 


inferior colliculus 
half of the midbrain tectum that is part of the brain stem auditory 
pathway 


inferior olive 
nucleus in the medulla that is involved in processing information 
related to motor control 


kinesthesia 
general sensory perception of movement of the body 


lateral column 
white matter of the spinal cord between the posterior horn on one side 
and the axons from the anterior horn on the same side; composed of 
many different groups of axons, of both ascending and descending 
tracts, carrying motor commands to and from the brain 


lateral hor 
region of the spinal cord gray matter in the thoracic, upper lumbar, and 
sacral regions that is the central component of the sympathetic division 
of the autonomic nervous system 


lateral sulcus 
surface landmark of the cerebral cortex that marks the boundary 
between the temporal lobe and the frontal and parietal lobes 


limbic cortex 
collection of structures of the cerebral cortex that are involved in 
emotion, memory, and behavior and are part of the larger limbic 
system 


limbic system 
structures at the edge (limit) of the boundary between the forebrain and 
hindbrain that are most associated with emotional behavior and 
memory formation 


longitudinal fissure 
large separation along the midline between the two cerebral 
hemispheres 


occipital lobe 
region of the cerebral cortex directly beneath the occipital bone of the 
cranium 


olfaction 


special sense responsible for smell, which has a unique, direct 
connection to the cerebrum 


parietal lobe 
region of the cerebral cortex directly beneath the parietal bone of the 
cranium 


parieto-occipital sulcus 
groove in the cerebral cortex representing the border between the 
parietal and occipital cortices 


postcentral gyrus 
primary motor cortex located in the frontal lobe of the cerebral cortex 


posterior columns 
white matter of the spinal cord that lies between the posterior horns of 
the gray matter, sometimes referred to as the dorsal column; composed 
of axons of ascending tracts that carry sensory information up to the 
brain 


posterior hor 
gray matter region of the spinal cord in which sensory input arrives, 
sometimes referred to as the dorsal horn 


posterior median sulcus 
midline feature of the posterior spinal cord, marking the separation 
between right and left sides of the cord 


posterolateral sulcus 
feature of the posterior spinal cord marking the entry of posterior nerve 
roots and the separation between the posterior and lateral columns of 
the white matter 


precentral gyrus 
ridge just posterior to the central sulcus, in the parietal lobe, where 


somatosensory processing initially takes place in the cerebrum 


prefrontal lobe 


specific region of the frontal lobe anterior to the more specific motor 
function areas, which can be related to the early planning of 
movements and intentions to the point of being personality-type 
functions 


premotor area 
region of the frontal lobe responsible for planning movements that will 
be executed through the primary motor cortex 


proprioception 
general sensory perceptions providing information about location and 
movement of body parts; the “sense of the self” 


putamen 
nucleus deep in the cerebrum that is part of the basal nuclei; along with 
the caudate, it is part of the striatum 


reticular formation 
diffuse region of gray matter throughout the brain stem that regulates 
Sleep, wakefulness, and states of consciousness 


somatosensation 
general senses related to the body, usually thought of as the senses of 
touch, which would include pain, temperature, and proprioception 


striatum 
the caudate and putamen collectively, as part of the basal nuclei, which 
receive input from the cerebral cortex 


subcortical nucleus 
all the nuclei beneath the cerebral cortex, including the basal nuclei 
and the basal forebrain 


substantia nigra pars compacta 
nuclei within the basal nuclei that release dopamine to modulate the 
function of the striatum; part of the motor pathway 


substantia nigra pars reticulata 


nuclei within the basal nuclei that serve as an output center of the 
nuclei; part of the motor pathway 


subthalamus 
nucleus within the basal nuclei that is part of the indirect pathway 


sulcus 
groove formed by convolutions in the surface of the cerebral cortex 


superior colliculus 
half of the midbrain tectum that is responsible for aligning visual, 
auditory, and somatosensory spatial perceptions 


tectum 
region of the midbrain, thought of as the roof of the cerebral aqueduct, 
which is subdivided into the inferior and superior colliculi 


tegmentum 
region of the midbrain, thought of as the floor of the cerebral aqueduct, 
which continues into the pons and medulla as the floor of the fourth 
ventricle 


temporal lobe 
region of the cerebral cortex directly beneath the temporal bone of the 
cranium 


thalamus 
major region of the diencephalon that is responsible for relaying 
information between the cerebrum and the hindbrain, spinal cord, and 
periphery 


ventral (anterior) nerve root 
axons emerging from the anterior or lateral horns of the spinal cord 


Circulation and the Central Nervous System 
By the end of this section, you will be able to: 


e Describe the vessels that supply the CNS with blood 

e Name the components of the ventricular system and the regions of the brain in which each is located 
e Explain the production of cerebrospinal fluid and its flow through the ventricles 

e Explain how a disruption in circulation would result in a stroke 


The CNS is crucial to the operation of the body, and any compromise in the brain and spinal cord can lead to 
severe difficulties. The CNS has a privileged blood supply, as suggested by the blood-brain barrier. The function of 
the tissue in the CNS is crucial to the survival of the organism, so the contents of the blood cannot simply pass into 
the central nervous tissue. To protect this region from the toxins and pathogens that may be traveling through the 
blood stream, there is strict control over what can move out of the general systems and into the brain and spinal 
cord. Because of this privilege, the CNS needs specialized structures for the maintenance of circulation. This 
begins with a unique arrangement of blood vessels carrying fresh blood into the CNS. Beyond the supply of blood, 
the CNS filters that blood into cerebrospinal fluid (CSF), which is then circulated through the cavities of the brain 
and spinal cord called ventricles. 


Blood Supply to the Brain 


A lack of oxygen to the CNS can be devastating, and the cardiovascular system has specific regulatory reflexes to 
ensure that the blood supply is not interrupted. There are multiple routes for blood to get into the CNS, with 
specializations to protect that blood supply and to maximize the ability of the brain to get an uninterrupted 
perfusion. 


Arterial Supply 


The major artery carrying recently oxygenated blood away from the heart is the aorta. The very first branches off 
the aorta supply the heart with nutrients and oxygen. The next branches give rise to the common carotid arteries, 
which further branch into the internal carotid arteries. The external carotid arteries supply blood to the tissues on 
the surface of the cranium. The bases of the common carotids contain stretch receptors that immediately respond to 
the drop in blood pressure upon standing. The orthostatic reflex is a reaction to this change in body position, so 
that blood pressure is maintained against the increasing effect of gravity (orthostatic means “standing up”). Heart 
rate increases—a reflex of the sympathetic division of the autonomic nervous system—and this raises blood 
pressure. 


The internal carotid artery enters the cranium through the carotid canal in the temporal bone. A second set of 
vessels that supply the CNS are the vertebral arteries, which are protected as they pass through the neck region 
by the transverse foramina of the cervical vertebrae. The vertebral arteries enter the cranium through the foramen 
magnum of the occipital bone. Branches off the left and right vertebral arteries merge into the anterior spinal 
artery supplying the anterior aspect of the spinal cord, found along the anterior median fissure. The two vertebral 
arteries then merge into the basilar artery, which gives rise to branches to the brain stem and cerebellum. The left 
and right internal carotid arteries and branches of the basilar artery all become the circle of Willis, a confluence of 
arteries that can maintain perfusion of the brain even if narrowing or a blockage limits flow through one part 
({link]). 

Circle of Willis 


Anterior cerebral 
Anterior artery 
communicating 


artery Ophthalmic 


artery 
Middle 


cerebral Anterior 


artery choroidal 
Internal artery 
carotid Posterior 
artery cerebral 


arte! 

Posterior me 
communicating 
artery 


Superior 
cerebellar 


artery 
Pontine ; 
arteries Basilar artery 
Anterior Vertebral 
inferior artery 


cerebellar 
artery . 
Posterior 
inferior 
cerebellar 
Anterior artely 
spinal artery 


The blood supply to the brain 
enters through the internal carotid 
arteries and the vertebral arteries, 
eventually giving rise to the circle 

of Willis. 


Watch this animation to see how blood flows to the brain and passes through the circle of Willis before being 
distributed through the cerebrum. The circle of Willis is a specialized arrangement of arteries that ensure constant 
perfusion of the cerebrum even in the event of a blockage of one of the arteries in the circle. The animation shows 
the normal direction of flow through the circle of Willis to the middle cerebral artery. Where would the blood 
come from if there were a blockage just posterior to the middle cerebral artery on the left? 


Venous Return 


After passing through the CNS, blood returns to the circulation through a series of dural sinuses and veins 
({link]). The superior sagittal sinus runs in the groove of the longitudinal fissure, where it absorbs CSF from the 
meninges. The superior sagittal sinus drains to the confluence of sinuses, along with the occipital sinuses and 
straight sinus, to then drain into the transverse sinuses. The transverse sinuses connect to the sigmoid sinuses, 


which then connect to the jugular veins. From there, the blood continues toward the heart to be pumped to the 
lungs for reoxygenation. 
Dural Sinuses and Veins 


Superior sagittal 
sinus 


Cranium Inferior sagittal 
Dura mater sinus 
Straight 
sinus 
Cerebral veins 
Tranverse 
sinus 
Great cerebral 


vein Confluence 


of sinuses 


Occipital 
sinus 


Blood returns to 
jugular vein via 
the sigmoid sinus 


Blood drains from the brain through a series of sinuses that 
connect to the jugular veins. 


Protective Coverings of the Brain and Spinal Cord 


The outer surface of the CNS is covered by a series of membranes composed of connective tissue called the 
meninges, which protect the brain. The dura mater is a thick fibrous layer and a strong protective sheath over the 
entire brain and spinal cord. It is anchored to the inner surface of the cranium and vertebral cavity. The arachnoid 
mater is a membrane of thin fibrous tissue that forms a loose sac around the CNS. Beneath the arachnoid is a thin, 
filamentous mesh called the arachnoid trabeculae, which looks like a spider web, giving this layer its name. 
Directly adjacent to the surface of the CNS is the pia mater, a thin fibrous membrane that follows the 
convolutions of gyri and sulci in the cerebral cortex and fits into other grooves and indentations ([link]). 
Meningeal Layers of Superior Sagittal Sinus 


Superior sagittal sinus 


Arachnoid mater Dura mater 
Subdural space 
Shee Arachnoid 


Pia mater granulation villi 


Arachnoid trabeculae Longitudinal fissure 


Cerebral cortex 


The layers of the meninges in the longitudinal fissure of 
the superior sagittal sinus are shown, with the dura mater 
adjacent to the inner surface of the cranium, the pia 
mater adjacent to the surface of the brain, and the 
arachnoid and subarachnoid space between them. An 


arachnoid villus is shown emerging into the dural sinus 
to allow CSF to filter back into the blood for drainage. 


Dura Mater 


Like a thick cap covering the brain, the dura mater is a tough outer covering. The name comes from the Latin for 
“tough mother” to represent its physically protective role. It encloses the entire CNS and the major blood vessels 
that enter the cranium and vertebral cavity. It is directly attached to the inner surface of the bones of the cranium 
and to the very end of the vertebral cavity. 


There are infoldings of the dura that fit into large crevasses of the brain. Two infoldings go through the midline 
separations of the cerebrum and cerebellum; one forms a shelf-like tent between the occipital lobes of the 
cerebrum and the cerebellum, and the other surrounds the pituitary gland. The dura also surrounds and supports the 
venous sinuses. 


Arachnoid Mater 


The middle layer of the meninges is the arachnoid, named for the spider-web-like trabeculae between it and the 
pia mater. The arachnoid defines a sac-like enclosure around the CNS. The trabeculae are found in the 
subarachnoid space, which is filled with circulating CSF. The arachnoid emerges into the dural sinuses as the 
arachnoid granulations, where the CSF is filtered back into the blood for drainage from the nervous system. 


The subarachnoid space is filled with circulating CSF, which also provides a liquid cushion to the brain and spinal 
cord. Similar to clinical blood work, a sample of CSF can be withdrawn to find chemical evidence of 
neuropathology or metabolic traces of the biochemical functions of nervous tissue. 


Pia Mater 


The outer surface of the CNS is covered in the thin fibrous membrane of the pia mater. It is thought to have a 
continuous layer of cells providing a fluid-impermeable membrane. The name pia mater comes from the Latin for 
“tender mother,” suggesting the thin membrane is a gentle covering for the brain. The pia extends into every 
convolution of the CNS, lining the inside of the sulci in the cerebral and cerebellar cortices. At the end of the 
spinal cord, a thin filament extends from the inferior end of CNS at the upper lumbar region of the vertebral 
column to the sacral end of the vertebral column. Because the spinal cord does not extend through the lower 
lumbar region of the vertebral column, a needle can be inserted through the dura and arachnoid layers to withdraw 
CSF. This procedure is called a lumbar puncture and avoids the risk of damaging the central tissue of the spinal 
cord. Blood vessels that are nourishing the central nervous tissue are between the pia mater and the nervous tissue. 


Note: 

Disorders of the... 

Meninges 

Meningitis is an inflammation of the meninges, the three layers of fibrous membrane that surround the CNS. 
Meningitis can be caused by infection by bacteria or viruses. The particular pathogens are not special to 
meningitis; it is just an inflammation of that specific set of tissues from what might be a broader infection. 
Bacterial meningitis can be caused by Streptococcus, Staphylococcus, or the tuberculosis pathogen, among many 
others. Viral meningitis is usually the result of common enteroviruses (such as those that cause intestinal 
disorders), but may be the result of the herpes virus or West Nile virus. Bacterial meningitis tends to be more 
severe. 


The symptoms associated with meningitis can be fever, chills, nausea, vomiting, light sensitivity, soreness of the 
neck, or severe headache. More important are the neurological symptoms, such as changes in mental state 
(confusion, memory deficits, and other dementia-type symptoms). A serious risk of meningitis can be damage to 
peripheral structures because of the nerves that pass through the meninges. Hearing loss is a common result of 
meningitis. 

The primary test for meningitis is a lumbar puncture. A needle inserted into the lumbar region of the spinal 
column through the dura mater and arachnoid membrane into the subarachnoid space can be used to withdraw the 
fluid for chemical testing. Fatality occurs in 5 to 40 percent of children and 20 to 50 percent of adults with 
bacterial meningitis. Treatment of bacterial meningitis is through antibiotics, but viral meningitis cannot be 
treated with antibiotics because viruses do not respond to that type of drug. Fortunately, the viral forms are milder. 


Note: 


ees 


= openstax couse 
mite 


Watch this video that describes the procedure known as the lumbar puncture, a medical procedure used to sample 
the CSF. Because of the anatomy of the CNS, it is a relative safe location to insert a needle. Why is the lumbar 
puncture performed in the lower lumbar area of the vertebral column? 


The Ventricular System 


Cerebrospinal fluid (CSF) circulates throughout and around the CNS. In other tissues, water and small molecules 
are filtered through capillaries as the major contributor to the interstitial fluid. In the brain, CSF is produced in 
special structures to perfuse through the nervous tissue of the CNS and is continuous with the interstitial fluid. 
Specifically, CSF circulates to remove metabolic wastes from the interstitial fluids of nervous tissues and return 
them to the blood stream. The ventricles are the open spaces within the brain where CSF circulates. In some of 
these spaces, CSF is produced by filtering of the blood that is performed by a specialized membrane known as a 
choroid plexus. The CSF circulates through all of the ventricles to eventually emerge into the subarachnoid space 
where it will be reabsorbed into the blood. 


The Ventricles 


There are four ventricles within the brain, all of which developed from the original hollow space within the neural 
tube, the central canal. The first two are named the lateral ventricles and are deep within the cerebrum. These 
ventricles are connected to the third ventricle by two openings called the interventricular foramina. The third 
ventricle is the space between the left and right sides of the diencephalon, which opens into the cerebral aqueduct 
that passes through the midbrain. The aqueduct opens into the fourth ventricle, which is the space between the 
cerebellum and the pons and upper medulla ([link]). 

Cerebrospinal Fluid Circulation 


Superior sagittal 
sinus 


Arachnoid granulation 


Subarachnoid space 


Choroid plexus Meningeal dura mater 


Right lateral ventricle 


Interventricular 
foramen 


Third ventricle 


Cerebral aqueduct 


AN : 
Lateral aperture SS Median aperture 


Fourth ventricle 
Central canal 


The choroid plexus in the four ventricles produce CSF, which 
is circulated through the ventricular system and then enters the 
subarachnoid space through the median and lateral apertures. 
The CSF is then reabsorbed into the blood at the arachnoid 
granulations, where the arachnoid membrane emerges into the 
dural sinuses. 


As the telencephalon enlarges and grows into the cranial cavity, it is limited by the space within the skull. The 
telencephalon is the most anterior region of what was the neural tube, but cannot grow past the limit of the frontal 
bone of the skull. Because the cerebrum fits into this space, it takes on a C-shaped formation, through the frontal, 
parietal, occipital, and finally temporal regions. The space within the telencephalon is stretched into this same C- 
shape. The two ventricles are in the left and right sides, and were at one time referred to as the first and second 
ventricles. The interventricular foramina connect the frontal region of the lateral ventricles with the third ventricle. 


The third ventricle is the space bounded by the medial walls of the hypothalamus and thalamus. The two thalami 
touch in the center in most brains as the massa intermedia, which is surrounded by the third ventricle. The cerebral 
aqueduct opens just inferior to the epithalamus and passes through the midbrain. The tectum and tegmentum of the 
midbrain are the roof and floor of the cerebral aqueduct, respectively. The aqueduct opens up into the fourth 
ventricle. The floor of the fourth ventricle is the dorsal surface of the pons and upper medulla (that gray matter 
making a continuation of the tegmentum of the midbrain). The fourth ventricle then narrows into the central canal 
of the spinal cord. 


The ventricular system opens up to the subarachnoid space from the fourth ventricle. The single median aperture 
and the pair of lateral apertures connect to the subarachnoid space so that CSF can flow through the ventricles 
and around the outside of the CNS. Cerebrospinal fluid is produced within the ventricles by a type of specialized 
membrane called a choroid plexus. Ependymal cells (one of the types of glial cells described in the introduction to 
the nervous system) surround blood capillaries and filter the blood to make CSF. The fluid is a clear solution with 
a limited amount of the constituents of blood. It is essentially water, small molecules, and electrolytes. Oxygen and 
carbon dioxide are dissolved into the CSF, as they are in blood, and can diffuse between the fluid and the nervous 
tissue. 


Cerebrospinal Fluid Circulation 


The choroid plexuses are found in all four ventricles. Observed in dissection, they appear as soft, fuzzy structures 
that may still be pink, depending on how well the circulatory system is cleared in preparation of the tissue. The 


CSF is produced from components extracted from the blood, so its flow out of the ventricles is tied to the pulse of 
cardiovascular circulation. 


From the lateral ventricles, the CSF flows into the third ventricle, where more CSF is produced, and then through 
the cerebral aqueduct into the fourth ventricle where even more CSF is produced. A very small amount of CSF is 
filtered at any one of the plexuses, for a total of about 500 milliliters daily, but it is continuously made and pulses 
through the ventricular system, keeping the fluid moving. From the fourth ventricle, CSF can continue down the 
central canal of the spinal cord, but this is essentially a cul-de-sac, so more of the fluid leaves the ventricular 
system and moves into the subarachnoid space through the median and lateral apertures. 


Within the subarachnoid space, the CSF flows around all of the CNS, providing two important functions. As with 
elsewhere in its circulation, the CSF picks up metabolic wastes from the nervous tissue and moves it out of the 
CNS. It also acts as a liquid cushion for the brain and spinal cord. By surrounding the entire system in the 
subarachnoid space, it provides a thin buffer around the organs within the strong, protective dura mater. The 
arachnoid granulations are outpocketings of the arachnoid membrane into the dural sinuses so that CSF can be 
reabsorbed into the blood, along with the metabolic wastes. From the dural sinuses, blood drains out of the head 
and neck through the jugular veins, along with the rest of the circulation for blood, to be reoxygenated by the lungs 
and wastes to be filtered out by the kidneys ((link]). 


Note: 


Watch this animation that shows the flow of CSF through the brain and spinal cord, and how it originates from the 
ventricles and then spreads into the space within the meninges, where the fluids then move into the venous sinuses 
to return to the cardiovascular circulation. What are the structures that produce CSF and where are they found? 
How are the structures indicated in this animation? 


Components of CSF Circulation 


Lateral Third Cerebral Fourth Central Subarachnoid 
ventricles ventricle aqueduct ventricle canal space 
Between 
: pons/upper : 
Location : . 7 Spinal External to 
in CNS Cerebrum Diencephalon Midbrain aes cond entire CNS 
cerebellum 
Blood Choroid Choroid Choroid Arachnoid 
vessel None None : 
plexus plexus plexus granulations 


structure 


Note: 

Disorders of the... 

Central Nervous System 

The supply of blood to the brain is crucial to its ability to perform many functions. Without a steady supply of 
oxygen, and to a lesser extent glucose, the nervous tissue in the brain cannot keep up its extensive electrical 
activity. These nutrients get into the brain through the blood, and if blood flow is interrupted, neurological 
function is compromised. 

The common name for a disruption of blood supply to the brain is a stroke. It is caused by a blockage to an artery 
in the brain. The blockage is from some type of embolus: a blood clot, a fat embolus, or an air bubble. When the 
blood cannot travel through the artery, the surrounding tissue that is deprived starves and dies. Strokes will often 
result in the loss of very specific functions. A stroke in the lateral medulla, for example, can cause a loss in the 
ability to swallow. Sometimes, seemingly unrelated functions will be lost because they are dependent on 
structures in the same region. Along with the swallowing in the previous example, a stroke in that region could 
affect sensory functions from the face or extremities because important white matter pathways also pass through 
the lateral medulla. Loss of blood flow to specific regions of the cortex can lead to the loss of specific higher 
functions, from the ability to recognize faces to the ability to move a particular region of the body. Severe or 
limited memory loss can be the result of a temporal lobe stroke. 

Related to strokes are transient ischemic attacks (TIAs), which can also be called “mini-strokes.” These are events 
in which a physical blockage may be temporary, cutting off the blood supply and oxygen to a region, but not to 
the extent that it causes cell death in that region. While the neurons in that area are recovering from the event, 
neurological function may be lost. Function can return if the area is able to recover from the event. 

Recovery from a stroke (or TIA) is strongly dependent on the speed of treatment. Often, the person who is present 
and notices something is wrong must then make a decision. The mnemonic FAST helps people remember what to 
look for when someone is dealing with sudden losses of neurological function. If someone complains of feeling 
“funny,” check these things quickly: Look at the person’s face. Does he or she have problems moving Face 
muscles and making regular facial expressions? Ask the person to raise his or her Arms above the head. Can the 
person lift one arm but not the other? Has the person’s Speech changed? Is he or she slurring words or having 
trouble saying things? If any of these things have happened, then it is ‘Time to call for help. 

Sometimes, treatment with blood-thinning drugs can alleviate the problem, and recovery is possible. If the tissue 
is damaged, the amazing thing about the nervous system is that it is adaptable. With physical, occupational, and 
speech therapy, victims of strokes can recover, or more accurately relearn, functions. 


Chapter Review 


The CNS has a privileged blood supply established by the blood-brain barrier. Establishing this barrier are 
anatomical structures that help to protect and isolate the CNS. The arterial blood to the brain comes from the 
internal carotid and vertebral arteries, which both contribute to the unique circle of Willis that provides constant 
perfusion of the brain even if one of the blood vessels is blocked or narrowed. That blood is eventually filtered to 
make a separate medium, the CSF, that circulates within the spaces of the brain and then into the surrounding 
space defined by the meninges, the protective covering of the brain and spinal cord. 


The blood that nourishes the brain and spinal cord is behind the glial-cell-enforced blood-brain barrier, which 
limits the exchange of material from blood vessels with the interstitial fluid of the nervous tissue. Thus, metabolic 
wastes are collected in cerebrospinal fluid that circulates through the CNS. This fluid is produced by filtering 
blood at the choroid plexuses in the four ventricles of the brain. It then circulates through the ventricles and into 
the subarachnoid space, between the pia mater and the arachnoid mater. From the arachnoid granulations, CSF is 
reabsorbed into the blood, removing the waste from the privileged central nervous tissue. 


The blood, now with the reabsorbed CSF, drains out of the cranium through the dural sinuses. The dura mater is 
the tough outer covering of the CNS, which is anchored to the inner surface of the cranial and vertebral cavities. It 
surrounds the venous space known as the dural sinuses, which connect to the jugular veins, where blood drains 
from the head and neck. 


Interactive Link Questions 


Exercise: 
Problem: 
Watch this animation to see how blood flows to the brain and passes through the circle of Willis before being 
distributed through the cerebrum. The circle of Willis is a specialized arrangement of arteries that ensure 
constant perfusion of the cerebrum even in the event of a blockage of one of the arteries in the circle. The 


animation shows the normal direction of flow through the circle of Willis to the middle cerebral artery. Where 
would the blood come from if there were a blockage just posterior to the middle cerebral artery on the left? 


Solution: 


If blood could not get to the middle cerebral artery through the posterior circulation, the blood would flow 
around the circle of Willis to reach that artery from an anterior vessel. Blood flow would just reverse within 
the circle. 


Exercise: 
Problem: 
Watch this video that describes the procedure known as the lumbar puncture, a medical procedure used to 


sample the CSF. Because of the anatomy of the CNS, it is a relative safe location to insert a needle. Why is 
the lumbar puncture performed in the lower lumbar area of the vertebral column? 


Solution: 


The spinal cord ends in the upper lumbar area of the vertebral column, so a needle inserted lower than that 
will not damage the nervous tissue of the CNS. 


Exercise: 
Problem: 
Watch this animation that shows the flow of CSF through the brain and spinal cord, and how it originates 
from the ventricles and then spreads into the space within the meninges, where the fluids then move into the 
venous sinuses to return to the cardiovascular circulation. What are the structures that produce CSF and where 
are they found? How are the structures indicated in this animation? 
Solution: 
The choroid plexuses of the ventricles make CSF. As shown, there is a little of the blue color appearing in 
each ventricle that is joined by the color flowing from the other ventricles. 


Review Questions 


Exercise: 


Problem: What blood vessel enters the cranium to supply the brain with fresh, oxygenated blood? 


a. common carotid artery 
b. jugular vein 

c. internal carotid artery 
d. aorta 


Solution: 


C 


Exercise: 


Problem: 


Which layer of the meninges surrounds and supports the sinuses that form the route through which blood 
drains from the CNS? 


a. dura mater 

b. arachnoid mater 
c. subarachnoid 

d. pia mater 


Solution: 


A 


Exercise: 


Problem: What type of glial cell is responsible for filtering blood to produce CSF at the choroid plexus? 


a. ependymal cell 
b. astrocyte 

c. oligodendrocyte 
d. Schwann cell 


Solution: 


A 


Exercise: 


Problem: Which portion of the ventricular system is found within the diencephalon? 


a. lateral ventricles 
b. third ventricle 

c. cerebral aqueduct 
d. fourth ventricle 


Solution: 
B 
Exercise: 
Problem:What condition causes a stroke? 


a. inflammation of meninges 

b. lumbar puncture 

c. infection of cerebral spinal fluid 
d. disruption of blood to the brain 


Solution: 


D 


Critical Thinking Questions 


Exercise: 
Problem: 


Why can the circle of Willis maintain perfusion of the brain even if there is a blockage in one part of the 
structure? 


Solution: 

The structure is a circular connection of blood vessels, so that blood coming up from one of the arteries can 

flow in either direction around the circle and avoid any blockage or narrowing of the blood vessels. 
Exercise: 

Problem: 


Meningitis is an inflammation of the meninges that can have severe effects on neurological function. Why is 
infection of this structure potentially so dangerous? 


Solution: 


The nerves that connect the periphery to the CNS pass through these layers of tissue and can be damaged by 
that inflammation, causing a loss of important neurological functions. 


Glossary 


anterior spinal artery 
blood vessel from the merged branches of the vertebral arteries that runs along the anterior surface of the 
spinal cord 


arachnoid granulation 
outpocket of the arachnoid membrane into the dural sinuses that allows for reabsorption of CSF into the blood 


arachnoid mater 
middle layer of the meninges named for the spider-web-like trabeculae that extend between it and the pia 
mater 


arachnoid trabeculae 
filaments between the arachnoid and pia mater within the subarachnoid space 


basilar artery 
blood vessel from the merged vertebral arteries that runs along the dorsal surface of the brain stem 


carotid canal 
opening in the temporal bone through which the internal carotid artery enters the cranium 


central canal 
hollow space within the spinal cord that is the remnant of the center of the neural tube 


cerebral aqueduct 
connection of the ventricular system between the third and fourth ventricles located in the midbrain 


choroid plexus 
specialized structures containing ependymal cells lining blood capillaries that filter blood to produce CSF in 
the four ventricles of the brain 


circle of Willis 
unique anatomical arrangement of blood vessels around the base of the brain that maintains perfusion of 
blood into the brain even if one component of the structure is blocked or narrowed 


common carotid artery 
blood vessel that branches off the aorta (or the brachiocephalic artery on the right) and supplies blood to the 
head and neck 


dura mater 
tough, fibrous, outer layer of the meninges that is attached to the inner surface of the cranium and vertebral 
column and surrounds the entire CNS 


dural sinus 
any of the venous structures surrounding the brain, enclosed within the dura mater, which drain blood from 
the CNS to the common venous return of the jugular veins 


foramen magnum 
large opening in the occipital bone of the skull through which the spinal cord emerges and the vertebral 
arteries enter the cranium 


fourth ventricle 
the portion of the ventricular system that is in the region of the brain stem and opens into the subarachnoid 
space through the median and lateral apertures 


internal carotid artery 
branch from the common carotid artery that enters the cranium and supplies blood to the brain 


interventricular foramina 
openings between the lateral ventricles and third ventricle allowing for the passage of CSF 


jugular veins 
blood vessels that return “used” blood from the head and neck 


lateral apertures 
pair of openings from the fourth ventricle to the subarachnoid space on either side and between the medulla 
and cerebellum 


lateral ventricles 
portions of the ventricular system that are in the region of the cerebrum 


lumbar puncture 
procedure used to withdraw CSF from the lower lumbar region of the vertebral column that avoids the risk of 
damaging CNS tissue because the spinal cord ends at the upper lumbar vertebrae 


median aperture 
singular opening from the fourth ventricle into the subarachnoid space at the midline between the medulla and 
cerebellum 


meninges 
protective outer coverings of the CNS composed of connective tissue 


occipital sinuses 
dural sinuses along the edge of the occipital lobes of the cerebrum 


orthostatic reflex 
sympathetic function that maintains blood pressure when standing to offset the increased effect of gravity 


pia mater 


thin, innermost membrane of the meninges that directly covers the surface of the CNS 


sigmoid sinuses 
dural sinuses that drain directly into the jugular veins 


straight sinus 
dural sinus that drains blood from the deep center of the brain to collect with the other sinuses 


subarachnoid space 
space between the arachnoid mater and pia mater that contains CSF and the fibrous connections of the 
arachnoid trabeculae 


superior sagittal sinus 
dural sinus that runs along the top of the longitudinal fissure and drains blood from the majority of the outer 
cerebrum 


third ventricle 
portion of the ventricular system that is in the region of the diencephalon 


transverse sinuses 
dural sinuses that drain along either side of the occipital—-cerebellar space 


ventricles 
remnants of the hollow center of the neural tube that are spaces for cerebrospinal fluid to circulate through the 
brain 


vertebral arteries 
arteries that ascend along either side of the vertebral column through the transverse foramina of the cervical 
vertebrae and enter the cranium through the foramen magnum 


Nerves and ganglia 
By the end of this section, you will be able to: 


e Describe the structures found in the PNS 

e Distinguish between somatic and autonomic structures, including the special peripheral structures of the 
enteric nervous system 

e Name the twelve cranial nerves and explain the functions associated with each 

e Describe the sensory and motor components of spinal nerves and the plexuses that they pass through 


The PNS is not as contained as the CNS because it is defined as everything that is not the CNS. Some peripheral 
structures are incorporated into the other organs of the body. In describing the anatomy of the PNS, it is necessary 
to describe the common structures, the nerves and the ganglia, as they are found in various parts of the body. Many 
of the neural structures that are incorporated into other organs are features of the digestive system; these structures 
are known as the enteric nervous system and are a special subset of the PNS. 


Ganglia 


A ganglion is a group of neuron cell bodies in the periphery. Ganglia can be categorized, for the most part, as 
either sensory ganglia or autonomic ganglia, referring to their primary functions. The most common type of 
sensory ganglion is a dorsal (posterior) root ganglion. These ganglia are the cell bodies of neurons with axons 
that are sensory endings in the periphery, such as in the skin, and that extend into the CNS through the dorsal nerve 
root. The ganglion is an enlargement of the nerve root. Under microscopic inspection, it can be seen to include the 
cell bodies of the neurons, as well as bundles of fibers that are the posterior nerve root ([link]). The cells of the 
dorsal root ganglion are unipolar cells, classifying them by shape. Also, the small round nuclei of satellite cells can 
be seen surrounding—as if they were orbiting—the neuron cell bodies. 

Dorsal Root Ganglion 


Pas. na 


The cell bodies of sensory neurons, which are unipolar neurons by 
shape, are seen in this photomicrograph. Also, the fibrous region is 
composed of the axons of these neurons that are passing through 
the ganglion to be part of the dorsal nerve root (tissue source: 
canine). LM x 40. (Micrograph provided by the Regents of 
University of Michigan Medical School © 2012) 


Spinal Cord and Root Ganglion 


The slide includes both a cross-section of the lumbar spinal cord 
and a section of the dorsal root ganglion (see also [link]) (tissue 
source: canine). LM x 1600. (Micrograph provided by the Regents 
of University of Michigan Medical School © 2012) 


Note: 
a 
=> openstax coLLece 
L 
= q 
ogy ae 


View the University of Michigan WebScope to explore the tissue sample in greater detail. If you zoom in on the 
dorsal root ganglion, you can see smaller satellite glial cells surrounding the large cell bodies of the sensory 
neurons. From what structure do satellite cells derive during embryologic development? 


Another type of sensory ganglion is a cranial nerve ganglion. This is analogous to the dorsal root ganglion, 
except that it is associated with a cranial nerve instead of a spinal nerve. The roots of cranial nerves are within 
the cranium, whereas the ganglia are outside the skull. For example, the trigeminal ganglion is superficial to the 
temporal bone whereas its associated nerve is attached to the mid-pons region of the brain stem. The neurons of 
cranial nerve ganglia are also unipolar in shape with associated satellite cells. 


The other major category of ganglia are those of the autonomic nervous system, which is divided into the 
sympathetic and parasympathetic nervous systems. The sympathetic chain ganglia constitute a row of ganglia 
along the vertebral column that receive central input from the lateral horn of the thoracic and upper lumbar spinal 
cord. Superior to the chain ganglia are three paravertebral ganglia in the cervical region. Three other autonomic 
ganglia that are related to the sympathetic chain are the prevertebral ganglia, which are located outside of the 
chain but have similar functions. They are referred to as prevertebral because they are anterior to the vertebral 
column. The neurons of these autonomic ganglia are multipolar in shape, with dendrites radiating out around the 
cell body where synapses from the spinal cord neurons are made. The neurons of the chain, paravertebral, and 
prevertebral ganglia then project to organs in the head and neck, thoracic, abdominal, and pelvic cavities to 
regulate the sympathetic aspect of homeostatic mechanisms. 


Another group of autonomic ganglia are the terminal ganglia that receive input from cranial nerves or sacral 
spinal nerves and are responsible for regulating the parasympathetic aspect of homeostatic mechanisms. These two 


sets of ganglia, sympathetic and parasympathetic, often project to the same organs—one input from the chain 
ganglia and one input from a terminal ganglion—to regulate the overall function of an organ. For example, the 
heart receives two inputs such as these; one increases heart rate, and the other decreases it. The terminal ganglia 
that receive input from cranial nerves are found in the head and neck, as well as the thoracic and upper abdominal 
cavities, whereas the terminal ganglia that receive sacral input are in the lower abdominal and pelvic cavities. 


Terminal ganglia below the head and neck are often incorporated into the wall of the target organ as a plexus. A 
plexus, in a general sense, is a network of fibers or vessels. This can apply to nervous tissue (as in this instance) or 
structures containing blood vessels (such as a choroid plexus). For example, the enteric plexus is the extensive 
network of axons and neurons in the wall of the small and large intestines. The enteric plexus is actually part of the 
enteric nervous system, along with the gastric plexuses and the esophageal plexus. Though the enteric nervous 
system receives input originating from central neurons of the autonomic nervous system, it does not require CNS 
input to function. In fact, it operates independently to regulate the digestive system. 


Nerves 


Bundles of axons in the PNS are referred to as nerves. These structures in the periphery are different than the 
central counterpart, called a tract. Nerves are composed of more than just nervous tissue. They have connective 
tissues invested in their structure, as well as blood vessels supplying the tissues with nourishment. The outer 
surface of a nerve is a surrounding layer of fibrous connective tissue called the epineurium. Within the nerve, 
axons are further bundled into fascicles, which are each surrounded by their own layer of fibrous connective tissue 
called perineurium. Finally, individual axons are surrounded by loose connective tissue called the endoneurium 
({link]). These three layers are similar to the connective tissue sheaths for muscles. Nerves are associated with the 
region of the CNS to which they are connected, either as cranial nerves connected to the brain or spinal nerves 
connected to the spinal cord. 

Nerve Structure 


Spinal nerve 


Epineurium 


Perineurium 


Blood vessels 
Axion 


Perineurium 


Endoneurium 


Perineurium 


Fascicles 


(b) 


The structure of a nerve is organized by the layers of 
connective tissue on the outside, around each fascicle, 
and surrounding the individual nerve fibers (tissue 
source: simian). LM x 40. (Micrograph provided by 
the Regents of University of Michigan Medical School 


mana 


© 2U12) 


Close-Up of Nerve Trunk 
; Wp asf 
SST 


Zoom in on this slide of a nerve trunk to examine the 
endoneurium, perineurium, and epineurium in greater detail (tissue 
source: simian). LM x 1600. (Micrograph provided by the Regents 

of University of Michigan Medical School © 2012) 


Note: 


— openstax coLLece* 
View the University of Michigan WebScope to explore the tissue sample in greater detail. With what structures in 
a skeletal muscle are the endoneurium, perineurium, and epineurium comparable? 


Cranial Nerves 


The nerves attached to the brain are the cranial nerves, which are primarily responsible for the sensory and motor 
functions of the head and neck (one of these nerves targets organs in the thoracic and abdominal cavities as part of 
the parasympathetic nervous system). There are twelve cranial nerves, which are designated CNI through CNXII 
for “Cranial Nerve,” using Roman numerals for 1 through 12. They can be classified as sensory nerves, motor 
nerves, or a combination of both, meaning that the axons in these nerves originate out of sensory ganglia external 
to the cranium or motor nuclei within the brain stem. Sensory axons enter the brain to synapse in a nucleus. Motor 
axons connect to skeletal muscles of the head or neck. Three of the nerves are solely composed of sensory fibers; 
five are strictly motor; and the remaining four are mixed nerves. 


Learning the cranial nerves is a tradition in anatomy courses, and students have always used mnemonic devices to 
remember the nerve names. A traditional mnemonic is the rhyming couplet, “On Old Olympus’ Towering Tops/A 
Finn And German Viewed Some Hops,” in which the initial letter of each word corresponds to the initial letter in 
the name of each nerve. The names of the nerves have changed over the years to reflect current usage and more 
accurate naming. An exercise to help learn this sort of information is to generate a mnemonic using words that 
have personal significance. The names of the cranial nerves are listed in [link] along with a brief description of 


their function, their source (sensory ganglion or motor nucleus), and their target (sensory nucleus or skeletal 
muscle). They are listed here with a brief explanation of each nerve ((Link]). 


The olfactory nerve and optic nerve are responsible for the sense of smell and vision, respectively. The 
oculomotor nerve is responsible for eye movements by controlling four of the extraocular muscles. It is also 
responsible for lifting the upper eyelid when the eyes point up, and for pupillary constriction. The trochlear nerve 
and the abducens nerve are both responsible for eye movement, but do so by controlling different extraocular 
muscles. The trigeminal nerve is responsible for cutaneous sensations of the face and controlling the muscles of 
mastication. The facial nerve is responsible for the muscles involved in facial expressions, as well as part of the 
sense of taste and the production of saliva. The vestibulocochlear nerve is responsible for the senses of hearing 
and balance. The glossopharyngeal nerve is responsible for controlling muscles in the oral cavity and upper 
throat, as well as part of the sense of taste and the production of saliva. The vagus nerve is responsible for 
contributing to homeostatic control of the organs of the thoracic and upper abdominal cavities. The spinal 
accessory nerve is responsible for controlling the muscles of the neck, along with cervical spinal nerves. The 
hypoglossal nerve is responsible for controlling the muscles of the lower throat and tongue. 

The Cranial Nerves 


Oculomotor 


nerve III Optic nerve II 


Trochlear 
IV 
nerve Trigeminal nerve V 
Abducens 
nerve VI 


. Facial nerve VII 
Vestibulocochlear 


nerve VIII 
Glossopharyngeal 
nerve IX 


Hypoglossal 


nerve XII 
Vagus nerve X 


The anatomical arrangement of the roots of the cranial 
nerves observed from an inferior view of the brain. 


Three of the cranial nerves also contain autonomic fibers, and a fourth is almost purely a component of the 
autonomic system. The oculomotor, facial, and glossopharyngeal nerves contain fibers that contact autonomic 
ganglia. The oculomotor fibers initiate pupillary constriction, whereas the facial and glossopharyngeal fibers both 
initiate salivation. The vagus nerve primarily targets autonomic ganglia in the thoracic and upper abdominal 
cavities. 


Note: 


CRS i] 
* 
=> Snensae COLLEGE" 
Os Ba 
Visit this site to read about a man who wakes with a headache and a loss of vision. His regular doctor sent him to 
an ophthalmologist to address the vision loss. The ophthalmologist recognizes a greater problem and immediately 


sends him to the emergency room. Once there, the patient undergoes a large battery of tests, but a definite cause 
cannot be found. A specialist recognizes the problem as meningitis, but the question is what caused it originally. 


How can that be cured? The loss of vision comes from swelling around the optic nerve, which probably presented 
as a bulge on the inside of the eye. Why is swelling related to meningitis going to push on the optic nerve? 


Another important aspect of the cranial nerves that lends itself to a mnemonic is the functional role each nerve 
plays. The nerves fall into one of three basic groups. They are sensory, motor, or both (see [link]). The sentence, 
“Some Say Marry Money But My Brother Says Brains Beauty Matter More,” corresponds to the basic function of 
each nerve. The first, second, and eighth nerves are purely sensory: the olfactory (CNI), optic (CNII), and 
vestibulocochlear (CNVIID nerves. The three eye-movement nerves are all motor: the oculomotor (CNHI), 
trochlear (CNIV), and abducens (CNVI). The spinal accessory (CNXI) and hypoglossal (CNXII) nerves are also 
strictly motor. The remainder of the nerves contain both sensory and motor fibers. They are the trigeminal (CNV), 
facial (CNVID, glossopharyngeal (CNIX), and vagus (CNX) nerves. The nerves that convey both are often related 
to each other. The trigeminal and facial nerves both concern the face; one concerns the sensations and the other 
concems the muscle movements. The facial and glossopharyngeal nerves are both responsible for conveying 
gustatory, or taste, sensations as well as controlling salivary glands. The vagus nerve is involved in visceral 
responses to taste, namely the gag reflex. This is not an exhaustive list of what these combination nerves do, but 
there is a thread of relation between them. 


Cranial Nerves 


Function 
Mnemonic # Name (S/M/B) Central connection (nuclei) 
On I Olfactory Smell (S) Olfactory bulb 
Old II Optic Vision (S) Hypothalamus/thalamus/midbrain 


Eye movements 


Olympus’ Il Oculomotor (M) Oculomotor nucleus 
Towering IV Trochlear Re movements Trochlear nucleus 

. F Sensory/motor Trigeminal nuclei in the 
Ops ” oo — face (B) midbrain, pons, and medulla 
A VI Abducens oe pvcnese Abducens nucleus 


(M) 


Cranial Nerves 


Function 
Mnemonic # Name (S/M/B) Central connection (nuclei) 
Finn VII Facial Motor — face, Facial nucleus, solitary nucleus, 

Taste (B) superior salivatory nucleus 
aa VII Auditory Hearing/balance Cochlear nucleus, Vestibular 

(Vestibulocochlear) (S) nucleus/cerebellum 

Motor teat Solitary nucleus, inferior 
German Ix Glossopharyngeal salivatory nucleus, nucleus 

Taste (B) : 

ambiguus 

Motor/sensory — 
Viewed Xx Vagus viscera Medulla 

(autonomic) (B) 

. Motor — head : 

Some XI Spinal Accessory and neck (M) Spinal accessory nucleus 
Hops XI Hypoglossal Motor lower Hypoglossal nucleus 


throat (M) 


Spinal Nerves 


The nerves connected to the spinal cord are the spinal nerves. The arrangement of these nerves is much more 
regular than that of the cranial nerves. All of the spinal nerves are combined sensory and motor axons that separate 
into two nerve roots. The sensory axons enter the spinal cord as the dorsal nerve root. The motor fibers, both 
somatic and autonomic, emerge as the ventral nerve root. The dorsal root ganglion for each nerve is an 
enlargement of the spinal nerve. 


There are 31 spinal nerves, named for the level of the spinal cord at which each one emerges. There are eight pairs 
of cervical nerves designated C1 to C8, twelve thoracic nerves designated T1 to T12, five pairs of lumbar nerves 


designated L1 to LS, five pairs of sacral nerves designated S1 to S5, and one pair of coccygeal nerves. The nerves 
are numbered from the superior to inferior positions, and each emerges from the vertebral column through the 
intervertebral foramen at its level. The first nerve, C1, emerges between the first cervical vertebra and the occipital 
bone. The second nerve, C2, emerges between the first and second cervical vertebrae. The same occurs for C3 to 
C7, but C8 emerges between the seventh cervical vertebra and the first thoracic vertebra. For the thoracic and 
lumbar nerves, each one emerges between the vertebra that has the same designation and the next vertebra in the 
column. The sacral nerves emerge from the sacral foramina along the length of that unique vertebra. 


Spinal nerves extend outward from the vertebral column to enervate the periphery. The nerves in the periphery are 
not straight continuations of the spinal nerves, but rather the reorganization of the axons in those nerves to follow 
different courses. Axons from different spinal nerves will come together into a systemic nerve. This occurs at four 
places along the length of the vertebral column, each identified as a nerve plexus, whereas the other spinal nerves 
directly correspond to nerves at their respective levels. In this instance, the word plexus is used to describe 
networks of nerve fibers with no associated cell bodies. 


Of the four nerve plexuses, two are found at the cervical level, one at the lumbar level, and one at the sacral level 
({link]). The cervical plexus is composed of axons from spinal nerves C1 through C5 and branches into nerves in 
the posterior neck and head, as well as the phrenic nerve, which connects to the diaphragm at the base of the 
thoracic cavity. The other plexus from the cervical level is the brachial plexus. Spinal nerves C4 through T1 
reorganize through this plexus to give rise to the nerves of the arms, as the name brachial suggests. A large nerve 
from this plexus is the radial nerve from which the axillary nerve branches to go to the armpit region. The radial 
nerve continues through the arm and is paralleled by the ulnar nerve and the median nerve. The lumbar plexus 
arises from all the lumbar spinal nerves and gives rise to nerves enervating the pelvic region and the anterior leg. 
The femoral nerve is one of the major nerves from this plexus, which gives rise to the saphenous nerve as a 
branch that extends through the anterior lower leg. The sacral plexus comes from the lower lumbar nerves L4 and 
L5 and the sacral nerves S1 to S4. The most significant systemic nerve to come from this plexus is the sciatic 
nerve, which is a combination of the tibial nerve and the fibular nerve. The sciatic nerve extends across the hip 
joint and is most commonly associated with the condition sciatica, which is the result of compression or irritation 
of the nerve or any of the spinal nerves giving rise to it. 


These plexuses are described as arising from spinal nerves and giving rise to certain systemic nerves, but they 
contain fibers that serve sensory functions or fibers that serve motor functions. This means that some fibers extend 
from cutaneous or other peripheral sensory surfaces and send action potentials into the CNS. Those are axons of 
sensory neurons in the dorsal root ganglia that enter the spinal cord through the dorsal nerve root. Other fibers are 
the axons of motor neurons of the anterior horn of the spinal cord, which emerge in the ventral nerve root and send 
action potentials to cause skeletal muscles to contract in their target regions. For example, the radial nerve contains 
fibers of cutaneous sensation in the arm, as well as motor fibers that move muscles in the arm. 


Spinal nerves of the thoracic region, T2 through T11, are not part of the plexuses but rather emerge and give rise to 
the intercostal nerves found between the ribs, which articulate with the vertebrae surrounding the spinal nerve. 
Nerve Plexuses of the Body 


Cervical plexus 


Phrenic nerve 
Brachial plexus 


Axillary nerve 
Median nerve ———_— 


Radial nerve — b 
Ulnar nerve ; @) 


© 


a} 


Ce CTa/ ealgeeyrraee 


ey 


1 


f 


OQ 


Lumbar plexus 


Femoral nerve 
Obturator nerve 


Sacral plexus 


Common 
fibular nerve 


There are four main nerve plexuses in the human 
body. The cervical plexus supplies nerves to the 
posterior head and neck, as well as to the 
diaphragm. The brachial plexus supplies nerves 
to the arm. The lumbar plexus supplies nerves to 
the anterior leg. The sacral plexus supplies nerves 
to the posterior leg. 


Note: 

Aging and the... 

Nervous System 

Anosmia is the loss of the sense of smell. It is often the result of the olfactory nerve being severed, usually 
because of blunt force trauma to the head. The sensory neurons of the olfactory epithelium have a limited lifespan 
of approximately one to four months, and new ones are made on a regular basis. The new neurons extend their 
axons into the CNS by growing along the existing fibers of the olfactory nerve. The ability of these neurons to be 
replaced is lost with age. Age-related anosmia is not the result of impact trauma to the head, but rather a slow loss 
of the sensory neurons with no new neurons born to replace them. 

Smell is an important sense, especially for the enjoyment of food. There are only five tastes sensed by the tongue, 
and two of them are generally thought of as unpleasant tastes (sour and bitter). The rich sensory experience of 


food is the result of odor molecules associated with the food, both as food is moved into the mouth, and therefore 
passes under the nose, and when it is chewed and molecules are released to move up the pharynx into the 
posterior nasal cavity. Anosmia results in a loss of the enjoyment of food. 

As the replacement of olfactory neurons declines with age, anosmia can set in. Without the sense of smell, many 
sufferers complain of food tasting bland. Often, the only way to enjoy food is to add seasoning that can be sensed 
on the tongue, which usually means adding table salt. The problem with this solution, however, is that this 
increases sodium intake, which can lead to cardiovascular problems through water retention and the associated 
increase in blood pressure. 


Chapter Review 


The PNS is composed of the groups of neurons (ganglia) and bundles of axons (nerves) that are outside of the 
brain and spinal cord. Ganglia are of two types, sensory or autonomic. Sensory ganglia contain unipolar sensory 
neurons and are found on the dorsal root of all spinal nerves as well as associated with many of the cranial nerves. 
Autonomic ganglia are in the sympathetic chain, the associated paravertebral or prevertebral ganglia, or in terminal 
ganglia near or within the organs controlled by the autonomic nervous system. 


Nerves are classified as cranial nerves or spinal nerves on the basis of their connection to the brain or spinal cord, 
respectively. The twelve cranial nerves can be strictly sensory in function, strictly motor in function, or a 
combination of the two functions. Sensory fibers are axons of sensory ganglia that carry sensory information into 
the brain and target sensory nuclei. Motor fibers are axons of motor neurons in motor nuclei of the brain stem and 
target skeletal muscles of the head and neck. Spinal nerves are all mixed nerves with both sensory and motor 
fibers. Spinal nerves emerge from the spinal cord and reorganize through plexuses, which then give rise to 
systemic nerves. Thoracic spinal nerves are not part of any plexus, but give rise to the intercostal nerves directly. 


Interactive Link Questions 


Exercise: 


Problem: 


[link] If you zoom in on the DRG, you can see smaller satellite glial cells surrounding the large cell bodies of 
the sensory neurons. From what structure do satellite cells derive during embryologic development? 


Solution: 


[link] They derive from the neural crest. 
Exercise: 


Problem: 


[link] To what structures in a skeletal muscle are the endoneurium, perineurium, and epineurium comparable? 


Solution: 


[link] The endoneurium surrounding individual nerve fibers is comparable to the endomysium surrounding 
myofibrils, the perineurium bundling axons into fascicles is comparable to the perimysium bundling muscle 
fibers into fascicles, and the epineurium surrounding the whole nerve is comparable to the epimysium 
surrounding the muscle. 


Exercise: 


Problem: 


Visit this site to read about a man who wakes with a headache and a loss of vision. His regular doctor sent 
him to an ophthalmologist to address the vision loss. The ophthalmologist recognizes a greater problem and 
immediately sends him to the emergency room. Once there, the patient undergoes a large battery of tests, but 
a definite cause cannot be found. A specialist recognizes the problem as meningitis, but the question is what 
caused it originally. How can that be cured? The loss of vision comes from swelling around the optic nerve, 
which probably presented as a bulge on the inside of the eye. Why is swelling related to meningitis going to 
push on the optic nerve? 


Solution: 


The optic nerve enters the CNS in its projection from the eyes in the periphery, which means that it crosses 
through the meninges. Meningitis will include swelling of those protective layers of the CNS, resulting in 
pressure on the optic nerve, which can compromise vision. 


Review Questions 


Exercise: 


Problem: What type of ganglion contains neurons that control homeostatic mechanisms of the body? 


a. sensory ganglion 

b. dorsal root ganglion 
c. autonomic ganglion 
d. cranial nerve ganglion 


Solution: 


GC 


Exercise: 


Problem:Which ganglion is responsible for cutaneous sensations of the face? 


a. otic ganglion 

b. vestibular ganglion 
c. geniculate ganglion 
d. trigeminal ganglion 


Solution: 


D 


Exercise: 


Problem:What is the name for a bundle of axons within a nerve? 


a. fascicle 

b. tract 

c. nerve root 
d. epineurium 


Solution: 


A 


Exercise: 


Problem:Which cranial nerve does not control functions in the head and neck? 


a. olfactory 

b. trochlear 

c. glossopharyngeal 
d. vagus 


Solution: 
D 
Exercise: 
Problem: Which of these structures is not under direct control of the peripheral nervous system? 
a. trigeminal ganglion 
b. gastric plexus 


c. sympathetic chain ganglia 
d. cervical plexus 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem:Why are ganglia and nerves not surrounded by protective structures like the meninges of the CNS? 


Solution: 


The peripheral nervous tissues are out in the body, sometimes part of other organ systems. There is not a 
privileged blood supply like there is to the brain and spinal cord, so peripheral nervous tissues do not need the 
same sort of protections. 


Exercise: 
Problem: 
Testing for neurological function involves a series of tests of functions associated with the cranial nerves. 


What functions, and therefore which nerves, are being tested by asking a patient to follow the tip of a pen 
with their eyes? 


Solution: 


The contraction of extraocular muscles is being tested, which is the function of the oculomotor, trochlear, and 
abducens nerves. 


Glossary 


abducens nerve 
sixth cranial nerve; responsible for contraction of one of the extraocular muscles 


axillary nerve 
systemic nerve of the arm that arises from the brachial plexus 


brachial plexus 
nerve plexus associated with the lower cervical spinal nerves and first thoracic spinal nerve 


cervical plexus 
nerve plexus associated with the upper cervical spinal nerves 


cranial nerve 
one of twelve nerves connected to the brain that are responsible for sensory or motor functions of the head 
and neck 


cranial nerve ganglion 
sensory ganglion of cranial nerves 


dorsal (posterior) root ganglion 
sensory ganglion attached to the posterior nerve root of a spinal nerve 


endoneurium 
innermost layer of connective tissue that surrounds individual axons within a nerve 


enteric nervous system 
peripheral structures, namely ganglia and nerves, that are incorporated into the digestive system organs 


enteric plexus 
neuronal plexus in the wall of the intestines, which is part of the enteric nervous system 


epineurium 
outermost layer of connective tissue that surrounds an entire nerve 


esophageal plexus 
neuronal plexus in the wall of the esophagus that is part of the enteric nervous system 


extraocular muscles 
six skeletal muscles that control eye movement within the orbit 


facial nerve 
seventh cranial nerve; responsible for contraction of the facial muscles and for part of the sense of taste, as 
well as causing saliva production 


fascicle 
small bundles of nerve or muscle fibers enclosed by connective tissue 


femoral nerve 
systemic nerve of the anterior leg that arises from the lumbar plexus 


fibular nerve 
systemic nerve of the posterior leg that begins as part of the sciatic nerve 


gastric plexuses 
neuronal networks in the wall of the stomach that are part of the enteric nervous system 


glossopharyngeal nerve 
ninth cranial nerve; responsible for contraction of muscles in the tongue and throat and for part of the sense of 
taste, as well as causing saliva production 


hypoglossal nerve 
twelfth cranial nerve; responsible for contraction of muscles of the tongue 


intercostal nerve 
systemic nerve in the thoracic cavity that is found between two ribs 


lumbar plexus 
nerve plexus associated with the lumbar spinal nerves 


median nerve 
systemic nerve of the arm, located between the ulnar and radial nerves 


nerve plexus 
network of nerves without neuronal cell bodies included 


oculomotor nerve 
third cranial nerve; responsible for contraction of four of the extraocular muscles, the muscle in the upper 
eyelid, and pupillary constriction 


olfactory nerve 
first cranial nerve; responsible for the sense of smell 


optic nerve 
second cranial nerve; responsible for visual sensation 


paravertebral ganglia 
autonomic ganglia superior to the sympathetic chain ganglia 


perineurium 
layer of connective tissue surrounding fascicles within a nerve 


phrenic nerve 
systemic nerve from the cervical plexus that enervates the diaphragm 


plexus 
network of nerves or nervous tissue 


prevertebral ganglia 
autonomic ganglia that are anterior to the vertebral column and functionally related to the sympathetic chain 
ganglia 


radial nerve 
systemic nerve of the arm, the distal component of which is located near the radial bone 


sacral plexus 
nerve plexus associated with the lower lumbar and sacral spinal nerves 


saphenous nerve 
systemic nerve of the lower anterior leg that is a branch from the femoral nerve 


sciatic nerve 
systemic nerve from the sacral plexus that is a combination of the tibial and fibular nerves and extends across 
the hip joint and gluteal region into the upper posterior leg 


sciatica 
painful condition resulting from inflammation or compression of the sciatic nerve or any of the spinal nerves 
that contribute to it 


spinal accessory nerve 
eleventh cranial nerve; responsible for contraction of neck muscles 


spinal nerve 
one of 31 nerves connected to the spinal cord 


sympathetic chain ganglia 
autonomic ganglia in a chain along the anterolateral aspect of the vertebral column that are responsible for 
contributing to homeostatic mechanisms of the autonomic nervous system 


systemic nerve 
nerve in the periphery distal to a nerve plexus or spinal nerve 


terminal ganglion 
autonomic ganglia that are near or within the walls of organs that are responsible for contributing to 
homeostatic mechanisms of the autonomic nervous system 


tibial nerve 
systemic nerve of the posterior leg that begins as part of the sciatic nerve 


trigeminal ganglion 
sensory ganglion that contributes sensory fibers to the trigeminal nerve 


trigeminal nerve 
fifth cranial nerve; responsible for cutaneous sensation of the face and contraction of the muscles of 
mastication 


trochlear nerve 
fourth cranial nerve; responsible for contraction of one of the extraocular muscles 


ulnar nerve 
systemic nerve of the arm located close to the ulna, a bone of the forearm 


vagus nerve 
tenth cranial nerve; responsible for the autonomic control of organs in the thoracic and upper abdominal 
cavities 


vestibulocochlear nerve 
eighth cranial nerve; responsible for the sensations of hearing and balance 


Sensory Perception 
By the end of this section, you will be able to: 


e Describe different types of sensory receptors 

e Describe the structures responsible for the special senses of taste, smell, 
hearing, balance, and vision 

e Distinguish how different tastes are transduced 

e Describe the means of mechanoreception for hearing and balance 

e List the supporting structures around the eye and describe the structure of 
the eyeball 

e Describe the processes of phototransduction 


A major role of sensory receptors is to help us learn about the environment 
around us, or about the state of our internal environment. Stimuli from varying 
sources, and of different types, are received and changed into the 
electrochemical signals of the nervous system. This occurs when a stimulus 
changes the cell membrane potential of a sensory neuron. The stimulus causes 
the sensory cell to produce an action potential that is relayed into the central 
nervous system (CNS), where it is integrated with other sensory information— 
or sometimes higher cognitive functions—to become a conscious perception of 
that stimulus. The central integration may then lead to a motor response. 


Describing sensory function with the term sensation or perception is a 
deliberate distinction. Sensation is the activation of sensory receptor cells at the 
level of the stimulus. Perception is the central processing of sensory stimuli 
into a meaningful pattern. Perception is dependent on sensation, but not all 
sensations are perceived. Receptors are the cells or structures that detect 
sensations. A receptor cell is changed directly by a stimulus. A transmembrane 
protein receptor is a protein in the cell membrane that mediates a physiological 
change in a neuron, most often through the opening of ion channels or changes 
in the cell signaling processes. Transmembrane receptors are activated by 
chemicals called ligands. For example, a molecule in food can serve as a ligand 
for taste receptors. Other transmembrane proteins, which are not accurately 
called receptors, are sensitive to mechanical or thermal changes. Physical 
changes in these proteins increase ion flow across the membrane, and can 
generate an action potential or a graded potential in the sensory neurons. 


Sensory Receptors 


Stimuli in the environment activate specialized receptor cells in the peripheral 
nervous system. Different types of stimuli are sensed by different types of 
receptor cells. Receptor cells can be classified into types on the basis of three 
different criteria: cell type, position, and function. Receptors can be classified 
structurally on the basis of cell type and their position in relation to stimuli 
they sense. They can also be classified functionally on the basis of the 
transduction of stimuli, or how the mechanical stimulus, light, or chemical 
changed the cell membrane potential. 


Structural Receptor Types 


The cells that interpret information about the environment can be either (1) a 
neuron that has a free nerve ending, with dendrites embedded in tissue that 
would receive a sensation; (2) a neuron that has an encapsulated ending in 
which the sensory nerve endings are encapsulated in connective tissue that 
enhances their sensitivity; or (3) a specialized receptor cell, which has distinct 
structural components that interpret a specific type of stimulus ((link]). The 
pain and temperature receptors in the dermis of the skin are examples of 
neurons that have free nerve endings. Also located in the dermis of the skin are 
lamellated corpuscles, neurons with encapsulated nerve endings that respond to 
pressure and touch. The cells in the retina that respond to light stimuli are an 
example of a specialized receptor, a photoreceptor. 

Receptor Classification by Cell Type 


Free nerve endings 
(dendrites) 


Axon Cell body 
L esaseaacats a a a a 


(a) Neuron (receptor) with free nerve endings 


Dendrite 


Axon 


Dee keeeedreernrsts 


(b) Neuron (receptor) with encapsulated nerve endings 


Encapsulated 


nerve ending Bipolar cell 


(c) Specialized receptor cell 


Receptor cell types can be classified on the basis of 
their structure. Sensory neurons can have either (a) free 
nerve endings or (b) encapsulated endings. 


Photoreceptors in the eyes, such as rod cells, are 
examples of (c) specialized receptor cells. These cells 
release neurotransmitters onto a bipolar cell, which then 
synapses with the optic nerve neurons. 


Another way that receptors can be classified is based on their location relative 
to the stimuli. An exteroceptor is a receptor that is located near a stimulus in 
the external environment, such as the somatosensory receptors that are located 
in the skin. An interoceptor is one that interprets stimuli from internal organs 
and tissues, such as the receptors that sense the increase in blood pressure in 
the aorta or carotid sinus. Finally, a proprioceptor is a receptor located near a 
moving part of the body, such as a muscle, that interprets the positions of the 
tissues as they move. 


Functional Receptor Types 


A third classification of receptors is by how the receptor transduces stimuli into 
membrane potential changes. Stimuli are of three general types. Some stimuli 
are ions and macromolecules that affect transmembrane receptor proteins when 
these chemicals diffuse across the cell membrane. Some stimuli are physical 
variations in the environment that affect receptor cell membrane potentials. 
Other stimuli include the electromagnetic radiation from visible light. For 
humans, the only electromagnetic energy that is perceived by our eyes is 
visible light. Some other organisms have receptors that humans lack, such as 
the heat sensors of snakes, the ultraviolet light sensors of bees, or magnetic 
receptors in migratory birds. 


Receptor cells can be further categorized on the basis of the type of stimuli 
they transduce. Chemical stimuli can be interpreted by a chemoreceptor that 
interprets chemical stimuli, such as an object’s taste or smell. Osmoreceptors 
respond to solute concentrations of body fluids. Additionally, pain is primarily 
a chemical sense that interprets the presence of chemicals from tissue damage, 
or similar intense stimuli, through a nociceptor. Physical stimuli, such as 
pressure and vibration, as well as the sensation of sound and body position 
(balance), are interpreted through a mechanoreceptor. Another physical 


stimulus that has its own type of receptor is temperature, which is sensed 
through a thermoreceptor that is either sensitive to temperatures above (heat) 
or below (cold) normal body temperature. 


Sensory Modalities 


Ask anyone what the senses are, and they are likely to list the five major senses 
—taste, smell, touch, hearing, and sight. However, these are not all of the 
senses. The most obvious omission from this list is balance. Also, what is 
referred to simply as touch can be further subdivided into pressure, vibration, 
stretch, and hair-follicle position, on the basis of the type of mechanoreceptors 
that perceive these touch sensations. Other overlooked senses include 
temperature perception by thermoreceptors and pain perception by nociceptors. 


Within the realm of physiology, senses can be classified as either general or 
specific. A general sense is one that is distributed throughout the body and has 
receptor cells within the structures of other organs. Mechanoreceptors in the 
skin, muscles, or the walls of blood vessels are examples of this type. General 
senses often contribute to the sense of touch, as described above, or to 
proprioception (body movement) and kinesthesia (body movement), or to a 
visceral sense, which is most important to autonomic functions. A special 
sense is one that has a specific organ devoted to it, namely the eye, inner ear, 
tongue, or nose. 


Each of the senses is referred to as a sensory modality. Modality refers to the 
way that information is encoded, which is similar to the idea of transduction. 
The main sensory modalities can be described on the basis of how each is 
transduced. The chemical senses are taste and smell. The general sense that is 
usually referred to as touch includes chemical sensation in the form of 
nociception, or pain. Pressure, vibration, muscle stretch, and the movement of 
hair by an external stimulus, are all sensed by mechanoreceptors. Hearing and 
balance are also sensed by mechanoreceptors. Finally, vision involves the 
activation of photoreceptors. 


Listing all the different sensory modalities, which can number as many as 17, 
involves separating the five major senses into more specific categories, or 

submodalities, of the larger sense. An individual sensory modality represents 
the sensation of a specific type of stimulus. For example, the general sense of 


touch, which is known as somatosensation, can be separated into light 
pressure, deep pressure, vibration, itch, pain, temperature, or hair movement. 


Gustation (Taste) 


Only a few recognized submodalities exist within the sense of taste, or 
gustation. Until recently, only four tastes were recognized: sweet, salty, sour, 
and bitter. Research at the turn of the 20th century led to recognition of the 
fifth taste, umami, during the mid-1980s. Umamii is a Japanese word that 
means “delicious taste,” and is often translated to mean savory. Very recent 
research has suggested that there may also be a sixth taste for fats, or lipids. 


Gustation is the special sense associated with the tongue. The surface of the 
tongue, along with the rest of the oral cavity, is lined by a stratified squamous 
epithelium. Raised bumps called papillae (singular = papilla) contain the 
structures for gustatory transduction. There are four types of papillae, based on 
their appearance ([link]): circumvallate, foliate, filiform, and fungiform. 
Within the structure of the papillae are taste buds that contain specialized 
gustatory receptor cells for the transduction of taste stimuli. These receptor 
cells are sensitive to the chemicals contained within foods that are ingested, 
and they release neurotransmitters based on the amount of the chemical in the 
food. Neurotransmitters from the gustatory cells can activate sensory neurons 
in the facial, glossopharyngeal, and vagus cranial nerves. 

The Tongue 


Taste buds 


Circumvallate papilla 


Taste hairs Taste pore 


Fungiform papilla Filiform papilla Foliate papilla 


Basal cell Transitional cell 


Gustatory cell 


The tongue is covered with small bumps, called papillae, 
which contain taste buds that are sensitive to chemicals in 
ingested food or drink. Different types of papillae are found 
in different regions of the tongue. The taste buds contain 
specialized gustatory receptor cells that respond to chemical 
stimuli dissolved in the saliva. These receptor cells activate 
sensory neurons that are part of the facial and 
glossopharyngeal nerves. LM x 1600. (Micrograph 
provided by the Regents of University of Michigan Medical 
School © 2012) 


Salty taste is simply the perception of sodium ions (Na’) in the saliva. When 
you eat something salty, the salt crystals dissociate into the component ions 
Na’ and CI, which dissolve into the saliva in your mouth. The Na* 
concentration becomes high outside the gustatory cells, creating a strong 
concentration gradient that drives the diffusion of the ion into the cells. The 


entry of Na* into these cells results in the depolarization of the cell membrane 
and the generation of a receptor potential. 


Sour taste is the perception of H* concentration. Just as with sodium ions in 
salty flavors, these hydrogen ions enter the cell and trigger depolarization. Sour 
flavors are, essentially, the perception of acids in our food. Increasing 
hydrogen ion concentrations in the saliva (lowering saliva pH) triggers 
progressively stronger graded potentials in the gustatory cells. For example, 
orange juice—which contains citric acid—will taste sour because it has a pH 
value of approximately 3. Of course, it is often sweetened so that the sour taste 
is masked. 


The first two tastes (salty and sour) are triggered by the cations Na” and H’. 
The other tastes result from food molecules binding to a G protein-coupled 
receptor. A G protein signal transduction system ultimately leads to 
depolarization of the gustatory cell. The sweet taste is the sensitivity of 
gustatory cells to the presence of glucose dissolved in the saliva. Other 
monosaccharides such as fructose, or artificial sweeteners such as aspartame 
(NutraSweet™), saccharine, or sucralose (Splenda™) also activate the sweet 
receptors. The affinity for each of these molecules varies, and some will taste 
sweeter than glucose because they bind to the G protein-coupled receptor 
differently. 


Bitter taste is similar to sweet in that food molecules bind to G protein—coupled 
receptors. However, there are a number of different ways in which this can 
happen because there are a large diversity of bitter-tasting molecules. Some 
bitter molecules depolarize gustatory cells, whereas others hyperpolarize 
gustatory cells. Likewise, some bitter molecules increase G protein activation 
within the gustatory cells, whereas other bitter molecules decrease G protein 
activation. The specific response depends on which molecule is binding to the 
receptor. 


One major group of bitter-tasting molecules are alkaloids. Alkaloids are 
nitrogen containing molecules that are commonly found in bitter-tasting plant 
products, such as coffee, hops (in beer), tannins (in wine), tea, and aspirin. By 
containing toxic alkaloids, the plant is less susceptible to microbe infection and 
less attractive to herbivores. 


Therefore, the function of bitter taste may primarily be related to stimulating 
the gag reflex to avoid ingesting poisons. Because of this, many bitter foods 
that are normally ingested are often combined with a sweet component to make 
them more palatable (cream and sugar in coffee, for example). The highest 
concentration of bitter receptors appear to be in the posterior tongue, where a 
gag reflex could still spit out poisonous food. 


The taste known as umami is often referred to as the savory taste. Like sweet 
and bitter, it is based on the activation of G protein-coupled receptors by a 
specific molecule. The molecule that activates this receptor is the amino acid 
L-glutamate. Therefore, the umami flavor is often perceived while eating 
protein-rich foods. Not surprisingly, dishes that contain meat are often 
described as savory. 


Once the gustatory cells are activated by the taste molecules, they release 
neurotransmitters onto the dendrites of sensory neurons. These neurons are part 
of the facial and glossopharyngeal cranial nerves, as well as a component 
within the vagus nerve dedicated to the gag reflex. The facial nerve connects to 
taste buds in the anterior third of the tongue. The glossopharyngeal nerve 
connects to taste buds in the posterior two thirds of the tongue. The vagus 
nerve connects to taste buds in the extreme posterior of the tongue, verging on 
the pharynx, which are more sensitive to noxious stimuli such as bitterness. 


Note: 


openstax COLLEGE 


3 


Watch this video to learn about Dr. Danielle Reed of the Monell Chemical 
Senses Center in Philadelphia, Pennsylvania, who became interested in 
science at an early age because of her sensory experiences. She recognized 
that her sense of taste was unique compared with other people she knew. Now, 
she studies the genetic differences between people and their sensitivities to 
taste stimuli. In the video, there is a brief image of a person sticking out their 


tongue, which has been covered with a colored dye. This is how Dr. Reed is 
able to visualize and count papillae on the surface of the tongue. People fall 
into two groups known as “tasters” and “non-tasters” based on the density of 
papillae on their tongue, which also indicates the number of taste buds. Non- 
tasters can taste food, but they are not as sensitive to certain tastes, such as 
bitterness. Dr. Reed discovered that she is a non-taster, which explains why 
she perceived bitterness differently than other people she knew. Are you very 
sensitive to tastes? Can you see any similarities among the members of your 
family? 


Olfaction (Smell) 


Like taste, the sense of smell, or olfaction, is also responsive to chemical 
stimuli. The olfactory receptor neurons are located in a small region within the 
superior nasal cavity ({link]). This region is referred to as the olfactory 
epithelium and contains bipolar sensory neurons. Each olfactory sensory 
neuron has dendrites that extend from the apical surface of the epithelium into 
the mucus lining the cavity. As airborne molecules are inhaled through the 
nose, they pass over the olfactory epithelial region and dissolve into the mucus. 
These odorant molecules bind to proteins that keep them dissolved in the 
mucus and help transport them to the olfactory dendrites. The odorant—protein 
complex binds to a receptor protein within the cell membrane of an olfactory 
dendrite. These receptors are G protein—coupled, and will produce a graded 
membrane potential in the olfactory neurons. 


The axon of an olfactory neuron extends from the basal surface of the 
epithelium, through an olfactory foramen in the cribriform plate of the ethmoid 
bone, and into the brain. The group of axons called the olfactory tract connect 
to the olfactory bulb on the ventral surface of the frontal lobe. From there, the 
axons split to travel to several brain regions. Some travel to the cerebrum, 
specifically to the primary olfactory cortex that is located in the inferior and 
medial areas of the temporal lobe. Others project to structures within the limbic 
system and hypothalamus, where smells become associated with long-term 
memory and emotional responses. This is how certain smells trigger emotional 
memories, such as the smell of food associated with one’s birthplace. Smell is 
the one sensory modality that does not synapse in the thalamus before 
connecting to the cerebral cortex. This intimate connection between the 


olfactory system and the cerebral cortex is one reason why smell can be a 
potent trigger of memories and emotion. 


The nasal epithelium, including the olfactory cells, can be harmed by airborne 
toxic chemicals. Therefore, the olfactory neurons are regularly replaced within 
the nasal epithelium, after which the axons of the new neurons must find their 
appropriate connections in the olfactory bulb. These new axons grow along the 
axons that are already in place in the cranial nerve. 

The Olfactory System 


Olfactory tract 


Olfactory bulb Olfactory 


tract 


Mitral cells 


Olfactory Olfactory neurons 


epithelium 


Nasalconchae = : 
Ethmoid bone 


Path of 
inhaled air 
Filaments of 
olfactory nerve 
Connective tissue 


Olfactory gland 


Olfactory receptor 


Dendrite 
Olfactory 


cilia Mucus 


Path of inhaled air containing 
odorant molecules 


(a) Nasal cavity (b) Olfactory system 


(c) Olfactory epithelium 


(a) The olfactory system begins in the peripheral 
structures of the nasal cavity. (b) The olfactory receptor 
neurons are within the olfactory epithelium. (c) Axons of 
the olfactory receptor neurons project through the 
cribriform plate of the ethmoid bone and synapse with 
the neurons of the olfactory bulb (tissue source: simian). 
LM x 812. (Micrograph provided by the Regents of 


University of Michigan Medical School © 2012) 


Note: 

Disorders of the... 

Olfactory System: Anosmia 

Blunt force trauma to the face, such as that common in many car accidents, 
can lead to the loss of the olfactory nerve, and subsequently, loss of the sense 
of smell. This condition is known as anosmia. When the frontal lobe of the 
brain moves relative to the ethmoid bone, the olfactory tract axons may be 
sheared apart. Professional fighters often experience anosmia because of 
repeated trauma to face and head. In addition, certain pharmaceuticals, such as 
antibiotics, can cause anosmia by killing all the olfactory neurons at once. If 
no axons are in place within the olfactory nerve, then the axons from newly 
formed olfactory neurons have no guide to lead them to their connections 
within the olfactory bulb. There are temporary causes of anosmia, as well, 
such as those caused by inflammatory responses related to respiratory 
infections or allergies. 

Loss of the sense of smell can result in food tasting bland. A person with an 
impaired sense of smell may require additional spice and seasoning levels for 
food to be tasted. Anosmia may also be related to some presentations of mild 
depression, because the loss of enjoyment of food may lead to a general sense 
of despair. 

The ability of olfactory neurons to replace themselves decreases with age, 
leading to age-related anosmia. This explains why some elderly people salt 
their food more than younger people do. However, this increased sodium 
intake can increase blood volume and blood pressure, increasing the risk of 
cardiovascular diseases in the elderly. 


Audition (Hearing) 


Hearing, or audition, is the transduction of sound waves into a neural signal 
that is made possible by the structures of the ear ([link]). The large, fleshy 
structure on the lateral aspect of the head is known as the auricle. Some 


sources will also refer to this structure as the pinna, though that term is more 
appropriate for a structure that can be moved, such as the external ear of a cat. 
The C-shaped curves of the auricle direct sound waves toward the auditory 
canal. The canal enters the skull through the external auditory meatus of the 
temporal bone. At the end of the auditory canal is the tympanic membrane, or 
ear drum, which vibrates after it is struck by sound waves. The auricle, ear 
canal, and tympanic membrane are often referred to as the external ear. The 
middle ear consists of a space spanned by three small bones called the 
ossicles. The three ossicles are the malleus, incus, and stapes, which are Latin 
names that roughly translate to hammer, anvil, and stirrup. The malleus is 
attached to the tympanic membrane and articulates with the incus. The incus, 
in turn, articulates with the stapes. The stapes is then attached to the inner ear, 
where the sound waves will be transduced into a neural signal. The middle ear 
is connected to the pharynx through the Eustachian tube, which helps 
equilibrate air pressure across the tympanic membrane. The tube is normally 
closed but will pop open when the muscles of the pharynx contract during 
swallowing or yawning. 

Structures of the Ear 


Malleus —_Incus Stapes (attached to 


oval window) 


Vestibule 


Auricle 
Vestibular nerve 


Cochlear nerve 


Round window 


Ear canal 
Cochlea 


: Eustachian tube 
Tympanic 
membrane cavity 


External ear Middle ear Inner ear 


Tympanic 


The external ear contains the auricle, ear canal, 
and tympanic membrane. The middle ear 
contains the ossicles and is connected to the 
pharynx by the Eustachian tube. The inner ear 
contains the cochlea and vestibule, which are 
responsible for audition and equilibrium, 
respectively. 


The inner ear is often described as a bony labyrinth, as it is composed of a 
series of canals embedded within the temporal bone. It has two separate 
regions, the cochlea and the vestibule, which are responsible for hearing and 
balance, respectively. The neural signals from these two regions are relayed to 
the brain stem through separate fiber bundles. However, these two distinct 
bundles travel together from the inner ear to the brain stem as the 
vestibulocochlear nerve. Sound is transduced into neural signals within the 
cochlear region of the inner ear, which contains the sensory neurons of the 
spiral ganglia. These ganglia are located within the spiral-shaped cochlea of 
the inner ear. The cochlea is attached to the stapes through the oval window. 


The oval window is located at the beginning of a fluid-filled tube within the 
cochlea called the scala vestibuli. The scala vestibuli extends from the oval 
window, travelling above the cochlear duct, which is the central cavity of the 
cochlea that contains the sound-transducing neurons. At the uppermost tip of 
the cochlea, the scala vestibuli curves over the top of the cochlear duct. The 
fluid-filled tube, now called the scala tympani, returns to the base of the 
cochlea, this time travelling under the cochlear duct. The scala tympani ends at 
the round window, which is covered by a membrane that contains the fluid 
within the scala. As vibrations of the ossicles travel through the oval window, 
the fluid of the scala vestibuli and scala tympani moves in a wave-like motion. 
The frequency of the fluid waves match the frequencies of the sound waves 
({link]). The membrane covering the round window will bulge out or pucker in 
with the movement of the fluid within the scala tympani. 

Transmission of Sound Waves to Cochlea 


@ Tympanic membrane _—_@) Vibrations are 
vibrates in response amplified across 
to sound wave. ossicles. 


(0) Sound wave represents alternating 
areas of high and low pressure. 


139d) 


} 


\ 
\\ 


| | 


Wavelength 


wy , 


Ne Z 


oe Frequency of sound wave : , , - ; 
measured in Hz (cycles ® Vibrations against oval window set up standing 
per second) wave in fluid of vestibuli. 


Scala 

’ vestibuli 

ima ; Cochlear 
nf duct 


Organ of Corti 


Basilar 
membrane 


Scala tympani 


© Pressure bends the membrane of the 


cochlear duct at a point of maximum WENEVAVAVAAVE 
vibration for a given frequency, causing 


hair cells in the basilar membrane to Frequency of standing 
vibrate. wave is the same as 
sound wave 


A sound wave causes the tympanic membrane to vibrate. 
This vibration is amplified as it moves across the malleus, 
incus, and stapes. The amplified vibration is picked up by 
the oval window causing pressure waves in the fluid of the 

scala vestibuli and scala tympani. The complexity of the 
pressure waves is determined by the changes in amplitude 

and frequency of the sound waves entering the ear. 


A cross-sectional view of the cochlea shows that the scala vestibuli and scala 
tympani run along both sides of the cochlear duct ([link]). The cochlear duct 
contains several organs of Corti, which tranduce the wave motion of the two 
scala into neural signals. The organs of Corti lie on top of the basilar 
membrane, which is the side of the cochlear duct located between the organs 
of Corti and the scala tympani. As the fluid waves move through the scala 
vestibuli and scala tympani, the basilar membrane moves at a specific spot, 
depending on the frequency of the waves. Higher frequency waves move the 
region of the basilar membrane that is close to the base of the cochlea. Lower 
frequency waves move the region of the basilar membrane that is near the tip 
of the cochlea. 


Cross Section of the Cochlea 


Bony cochlear wall 


Scala vestibuli 
Cochlear duct 
Tectorial membrane 
Basilar membrane 


Scala tympani 


— ———— Cochlear branch 
of N VIII 


The three major spaces within the cochlea are 
highlighted. The scala tympani and scala vestibuli lie 
on either side of the cochlear duct. The organ of Corti, 
containing the mechanoreceptor hair cells, is adjacent 

to the scala tympani, where it sits atop the basilar 
membrane. 


The organs of Corti contain hair cells, which are named for the hair-like 
stereocilia extending from the cell’s apical surfaces ([{link]). The stereocilia are 
an array of microvilli-like structures arranged from tallest to shortest. Protein 
fibers tether adjacent hairs together within each array, such that the array will 
bend in response to movements of the basilar membrane. The stereocilia 
extend up from the hair cells to the overlying tectorial membrane, which is 
attached medially to the organ of Corti. When the pressure waves from the 
scala move the basilar membrane, the tectorial membrane slides across the 
stereocilia. This bends the stereocilia either toward or away from the tallest 
member of each array. When the stereocilia bend toward the tallest member of 
their array, tension in the protein tethers opens ion channels in the hair cell 
membrane. This will depolarize the hair cell membrane, triggering nerve 
impulses that travel down the afferent nerve fibers attached to the hair cells. 
When the stereocilia bend toward the shortest member of their array, the 
tension on the tethers slackens and the ion channels close. When no sound is 
present, and the stereocilia are standing straight, a small amount of tension still 


exists on the tethers, keeping the membrane potential of the hair cell slightly 
depolarized. 

Hair Cell 

ge tia 


Tether 


Stereocilia 


Hair cell 


The hair cell is a mechanoreceptor with an array of 
stereocilia emerging from its apical surface. The 
stereocilia are tethered together by proteins that open ion 
channels when the array is bent toward the tallest 
member of their array, and closed when the array is bent 
toward the shortest member of their array. 


Cochlea and Organ of Corti 


LM x 412. (Micrograph provided by the Regents 
of University of Michigan Medical School © 
2012) 


Note: 
ae 
— openstax COLLEGE 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. The basilar membrane is the thin membrane that extends from 
the central core of the cochlea to the edge. What is anchored to this membrane 
so that they can be activated by movement of the fluids within the cochlea? 


As stated above, a given region of the basilar membrane will only move if the 
incoming sound is at a specific frequency. Because the tectorial membrane 
only moves where the basilar membrane moves, the hair cells in this region 
will also only respond to sounds of this specific frequency. Therefore, as the 
frequency of a sound changes, different hair cells are activated all along the 
basilar membrane. The cochlea encodes auditory stimuli for frequencies 
between 20 and 20,000 Hz, which is the range of sound that human ears can 
detect. The unit of Hertz measures the frequency of sound waves in terms of 
cycles produced per second. Frequencies as low as 20 Hz are detected by hair 
cells at the apex, or tip, of the cochlea. Frequencies in the higher ranges of 20 
KHz are encoded by hair cells at the base of the cochlea, close to the round and 
oval windows ((link]). Most auditory stimuli contain a mixture of sounds at a 
variety of frequencies and intensities (represented by the amplitude of the 
sound wave). The hair cells along the length of the cochlear duct, which are 
each sensitive to a particular frequency, allow the cochlea to separate auditory 
stimuli by frequency, just as a prism separates visible light into its component 
colors. 

Frequency Coding in the Cochlea 


Oval window base 


Tectorial 
membrane 


20,000 Hz 1500 Hz 20 Hz 
(high frequency) = (medium frequency) (low frequency) 


_— 


Relative length of fibers in basilar membrane 


Basilar 
membrane 


Round window 


The standing sound wave generated in the cochlea by the 
movement of the oval window deflects the basilar 
membrane on the basis of the frequency of sound. 
Therefore, hair cells at the base of the cochlea are 

activated only by high frequencies, whereas those at the 
apex of the cochlea are activated only by low 
frequencies. 


Note: 


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pene Ea) 
_ openstax COLLEGE” 


. 7, 


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Watch this video to learn more about how the structures of the ear convert 
sound waves into a neural signal by moving the “hairs,” or stereocilia, of the 
cochlear duct. Specific locations along the length of the duct encode specific 
frequencies, or pitches. The brain interprets the meaning of the sounds we 


hear as music, speech, noise, etc. Which ear structures are responsible for the 
amplification and transfer of sound from the external ear to the inner ear? 


Note: 


mms Openstax COLLEGE 
| Te on Fa tae 


Watch this animation to learn more about the inner ear and to see the cochlea 
unroll, with the base at the back of the image and the apex at the front. 
Specific wavelengths of sound cause specific regions of the basilar membrane 
to vibrate, much like the keys of a piano produce sound at different 
frequencies. Based on the animation, where do frequencies—from high to low 
pitches—cause activity in the hair cells within the cochlear duct? 


Equilibrium (Balance) 


Along with audition, the inner ear is responsible for encoding information 
about equilibrium, the sense of balance. A similar mechanoreceptor—a hair 
cell with stereocilia—senses head position, head movement, and whether our 
bodies are in motion. These cells are located within the vestibule of the inner 
ear. Head position is sensed by the utricle and saccule, whereas head 
movement is sensed by the semicircular canals. The neural signals generated 
in the vestibular ganglion are transmitted through the vestibulocochlear nerve 
to the brain stem and cerebellum. 


The utricle and saccule are both largely composed of macula tissue (plural = 
maculae). The macula is composed of hair cells surrounded by support cells. 
The stereocilia of the hair cells extend into a viscous gel called the otolithic 
membrane ((link]). On top of the otolithic membrane is a layer of calcium 
carbonate crystals, called otoliths. The otoliths essentially make the otolithic 
membrane top-heavy. The otolithic membrane moves separately from the 


macula in response to head movements. Tilting the head causes the otolithic 
membrane to slide over the macula in the direction of gravity. The moving 
otolithic membrane, in turn, bends the sterocilia, causing some hair cells to 
depolarize as others hyperpolarize. The exact position of the head is interpreted 
by the brain based on the pattern of hair-cell depolarization. 

Linear Acceleration Coding by Maculae 


Otolithic 


Otoliths 
membrane i 
2 a Teh = 2-5 


Head upright 
Endolymph 
Macula 


Otoliths 


Otolithic 
membrane 


Hair cells 


Vestibular division of ‘ - 
vestibulocochlear nerve Head tilted forward 


aC 


\ 


The maculae are specialized for sensing linear acceleration, 
such as when gravity acts on the tilting head, or if the head 
starts moving in a straight line. The difference in inertia 
between the hair cell stereocilia and the otolithic membrane 
in which they are embedded leads to a shearing force that 
causes the stereocilia to bend in the direction of that linear 
acceleration. 


The semicircular canals are three ring-like extensions of the vestibule. One is 
oriented in the horizontal plane, whereas the other two are oriented in the 
vertical plane. The anterior and posterior vertical canals are oriented at 
approximately 45 degrees relative to the sagittal plane ([link]). The base of 
each semicircular canal, where it meets with the vestibule, connects to an 
enlarged region known as the ampulla. The ampulla contains the hair cells that 
respond to rotational movement, such as turning the head while saying “no.” 
The stereocilia of these hair cells extend into the cupula, a membrane that 
attaches to the top of the ampulla. As the head rotates in a plane parallel to the 
semicircular canal, the fluid lags, deflecting the cupula in the direction 


opposite to the head movement. The semicircular canals contain several 
ampullae, with some oriented horizontally and others oriented vertically. By 
comparing the relative movements of both the horizontal and vertical 
ampullae, the vestibular system can detect the direction of most head 
movements within three-dimensional (3-D) space. 

Rotational Coding by Semicircular Canals 


h As the head rotates, 
Ampullary nerve \\\\\) cupula bends in opposite 
direction of the rotation 


Rotational movement of the head is encoded by the hair 
cells in the base of the semicircular canals. As one of the 
canals moves in an arc with the head, the internal fluid 
moves in the opposite direction, causing the cupula and 
stereocilia to bend. The movement of two canals within a 
plane results in information about the direction in which the 
head is moving, and activation of all six canals can give a 
very precise indication of head movement in three 
dimensions. 


Somatosensation (Touch) 


Somatosensation is considered a general sense, as opposed to the special senses 
discussed in this section. Somatosensation is the group of sensory modalities 
that are associated with touch, proprioception, and interoception. These 
modalities include pressure, vibration, light touch, tickle, itch, temperature, 
pain, proprioception, and kinesthesia. This means that its receptors are not 


associated with a specialized organ, but are instead spread throughout the body 
in a variety of organs. Many of the somatosensory receptors are located in the 
skin, but receptors are also found in muscles, tendons, joint capsules, 
ligaments, and in the walls of visceral organs. 


Two types of somatosensory signals that are transduced by free nerve endings 
are pain and temperature. These two modalities use thermoreceptors and 
nociceptors to transduce temperature and pain stimuli, respectively. 
Temperature receptors are stimulated when local temperatures differ from body 
temperature. Some thermoreceptors are sensitive to just cold and others to just 
heat. Nociception is the sensation of potentially damaging stimuli. Mechanical, 
chemical, or thermal stimuli beyond a set threshold will elicit painful 
sensations. Stressed or damaged tissues release chemicals that activate receptor 
proteins in the nociceptors. For example, the sensation of heat associated with 
spicy foods involves capsaicin, the active molecule in hot peppers. Capsaicin 
molecules bind to a transmembrane ion channel in nociceptors that is sensitive 
to temperatures above 37°C. The dynamics of capsaicin binding with this 
transmembrane ion channel is unusual in that the molecule remains bound for a 
long time. Because of this, it will decrease the ability of other stimuli to elicit 
pain sensations through the activated nociceptor. For this reason, capsaicin can 
be used as a topical analgesic, such as in products such as Icy Hot™. 


If you drag your finger across a textured surface, the skin of your finger will 
vibrate. Such low frequency vibrations are sensed by mechanoreceptors called 
Merkel cells, also known as type I cutaneous mechanoreceptors. Merkel cells 
are located in the stratum basale of the epidermis. Deep pressure and vibration 
is transduced by lamellated (Pacinian) corpuscles, which are receptors with 
encapsulated endings found deep in the dermis, or subcutaneous tissue. Light 
touch is transduced by the encapsulated endings known as tactile (Meissner) 
corpuscles. Follicles are also wrapped in a plexus of nerve endings known as 
the hair follicle plexus. These nerve endings detect the movement of hair at the 
surface of the skin, such as when an insect may be walking along the skin. 
Stretching of the skin is transduced by stretch receptors known as bulbous 
corpuscles. Bulbous corpuscles are also known as Ruffini corpuscles, or type II 
cutaneous mechanoreceptors. 


Other somatosensory receptors are found in the joints and muscles. Stretch 
receptors monitor the stretching of tendons, muscles, and the components of 
joints. For example, have you ever stretched your muscles before or after 


exercise and noticed that you can only stretch so far before your muscles 
spasm back to a less stretched state? This spasm is a reflex that is initiated by 
stretch receptors to avoid muscle tearing. Such stretch receptors can also 
prevent over-contraction of a muscle. In skeletal muscle tissue, these stretch 
receptors are called muscle spindles. Golgi tendon organs similarly transduce 
the stretch levels of tendons. Bulbous corpuscles are also present in joint 
capsules, where they measure stretch in the components of the skeletal system 
within the joint. The types of nerve endings, their locations, and the stimuli 
they transduce are presented in [link]. 


Mechanoreceptors of Somatosensation 


Name 


Free nerve 
endings 


Mechanoreceptors 


Bulbous 
corpuscle 


Historical 
(eponymous) 
name 


Merkel’s 
discs 


Ruffini’s 
corpuscle 


Location(s) 


Dermis, 
cornea, 
tongue, joint 
capsules, 
visceral 
organs 


Epidermal— 
dermal 
junction, 
mucosal 
membranes 


Dermis, joint 
capsules 


Stimuli 


Pain, 
temperature, 
mechanical 
deformation 


Low 
frequency 
vibration 
(5-15 Hz) 


Stretch 


Mechanoreceptors of Somatosensation 


Name 


Tactile corpuscle 


Lamellated 
corpuscle 


Hair follicle 
plexus 


Muscle spindle 


Tendon stretch 
organ 


Historical 
(eponymous) 
name 


Meissner’s 


corpuscle 


Pacinian 
corpuscle 


Golgi tendon 
organ 


*No corresponding eponymous name. 


Vision 


Location(s) 


Papillary 
dermis, 
especially in 
the fingertips 
and lips 


Deep dermis, 
subcutaneous 
tissue 


Wrapped 
around hair 
follicles in 
the dermis 


In line with 
skeletal 
muscle fibers 


In line with 
tendons 


Stimuli 


Light touch, 
vibrations 
below 50 
Hz 


Deep 
pressure, 
high- 
frequency 
vibration 
(around 250 
Hz) 


Movement 
of hair 


Muscle 
contraction 
and stretch 


Stretch of 
tendons 


Vision is the special sense of sight that is based on the transduction of light 
stimuli received through the eyes. The eyes are located within either orbit in 
the skull. The bony orbits surround the eyeballs, protecting them and anchoring 
the soft tissues of the eye ({link]). The eyelids, with lashes at their leading 
edges, help to protect the eye from abrasions by blocking particles that may 
land on the surface of the eye. The inner surface of each lid is a thin membrane 
known as the palpebral conjunctiva. The conjunctiva extends over the white 
areas of the eye (the sclera), connecting the eyelids to the eyeball. Tears are 
produced by the lacrimal gland, located beneath the lateral edges of the nose. 
Tears produced by this gland flow through the lacrimal duct to the medial 
corner of the eye, where the tears flow over the conjunctiva, washing away 
foreign particles. 


The Eye in the Orbit 
ign _— ; Eyebrow 


i 
we Aas Orbicularis oculi 


muscle 


Levator palpebrae 
superioris muscle 


Palpebral conjunctiva 


Eyelashes 


Cornea 


Conjunctiva 


The eye is located within the orbit and 
surrounded by soft tissues that protect and 
support its function. The orbit is surrounded by 
cranial bones of the skull. 


Movement of the eye within the orbit is accomplished by the contraction of six 
extraocular muscles that originate from the bones of the orbit and insert into 
the surface of the eyeball ([link]). Four of the muscles are arranged at the 
cardinal points around the eye and are named for those locations. They are the 
superior rectus, medial rectus, inferior rectus, and lateral rectus. When 


each of these muscles contract, the eye to moves toward the contracting 
muscle. For example, when the superior rectus contracts, the eye rotates to 
look up. The superior oblique originates at the posterior orbit, near the origin 
of the four rectus muscles. However, the tendon of the oblique muscles threads 
through a pulley-like piece of cartilage known as the trochlea. The tendon 
inserts obliquely into the superior surface of the eye. The angle of the tendon 
through the trochlea means that contraction of the superior oblique rotates the 
eye medially. The inferior oblique muscle originates from the floor of the 
orbit and inserts into the inferolateral surface of the eye. When it contracts, it 
laterally rotates the eye, in opposition to the superior oblique. Rotation of the 
eye by the two oblique muscles is necessary because the eye is not perfectly 
aligned on the sagittal plane. When the eye looks up or down, the eye must 
also rotate slightly to compensate for the superior rectus pulling at 
approximately a 20-degree angle, rather than straight up. The same is true for 
the inferior rectus, which is compensated by contraction of the inferior oblique. 
A seventh muscle in the orbit is the levator palpebrae superioris, which is 
responsible for elevating and retracting the upper eyelid, a movement that 
usually occurs in concert with elevation of the eye by the superior rectus (see 
[link]). 


The extraocular muscles are innervated by three cranial nerves. The lateral 
rectus, which causes abduction of the eye, is innervated by the abducens nerve. 
The superior oblique is innervated by the trochlear nerve. All of the other 
muscles are innervated by the oculomotor nerve, as is the levator palpebrae 
superioris. The motor nuclei of these cranial nerves connect to the brain stem, 
which coordinates eye movements. 

Extraocular Muscles 


Superior oblique 
muscle 


Trochlea Superior x Trochlea 
; ‘ rectus ae 
Superior oblique 
tendon Superior 
Superior rectus oblique 
muscle 
Lateral rectus 
muscle r =; 
Lateral rectus a Medial rectus 
Inferior Inferior 
oblique rectus 


Common Inferior rectus Inferior oblique SS 
tendinous ring muscle muscle 


Lateral view of the right eye Anterior view of the right eye 


The extraocular muscles move the eye within the orbit. 


The eye itself is a hollow sphere composed of three layers of tissue. The 
outermost layer is the fibrous tunic, which includes the white sclera and clear 
cornea. The sclera accounts for five sixths of the surface of the eye, most of 
which is not visible, though humans are unique compared with many other 
species in having so much of the “white of the eye” visible ({link]). The 
transparent cornea covers the anterior tip of the eye and allows light to enter 
the eye. The middle layer of the eye is the vascular tunic, which is mostly 
composed of the choroid, ciliary body, and iris. The choroid is a layer of 
highly vascularized connective tissue that provides a blood supply to the 
eyeball. The choroid is posterior to the ciliary body, a muscular structure that 
is attached to the lens by suspensory ligaments, or zonule fibers. These two 
structures bend the lens, allowing it to focus light on the back of the eye. 
Overlaying the ciliary body, and visible in the anterior eye, is the iris—the 
colored part of the eye. The iris is a smooth muscle that opens or closes the 
pupil, which is the hole at the center of the eye that allows light to enter. The 
iris constricts the pupil in response to bright light and dilates the pupil in 
response to dim light. The innermost layer of the eye is the neural tunic, or 
retina, which contains the nervous tissue responsible for photoreception. 


The eye is also divided into two cavities: the anterior cavity and the posterior 
cavity. The anterior cavity is the space between the cornea and lens, including 
the iris and ciliary body. It is filled with a watery fluid called the aqueous 
humor. The posterior cavity is the space behind the lens that extends to the 
posterior side of the interior eyeball, where the retina is located. The posterior 
cavity is filled with a more viscous fluid called the vitreous humor. 


The retina is composed of several layers and contains specialized cells for the 
initial processing of visual stimuli. The photoreceptors (rods and cones) change 
their membrane potential when stimulated by light energy. The change in 
membrane potential alters the amount of neurotransmitter that the 
photoreceptor cells release onto bipolar cells in the outer synaptic layer. It is 
the bipolar cell in the retina that connects a photoreceptor to a retinal ganglion 
cell (RGC) in the inner synaptic layer. There, amacrine cells additionally 
contribute to retinal processing before an action potential is produced by the 
RGC. The axons of RGCs, which lie at the innermost layer of the retina, 
collect at the optic disc and leave the eye as the optic nerve (see [link]). 
Because these axons pass through the retina, there are no photoreceptors at the 


very back of the eye, where the optic nerve begins. This creates a “blind spot” 
in the retina, and a corresponding blind spot in our visual field. 
Structure of the Eye 


Lateral 


Posterior cavity 
Vitreous chamber 


; N 


Scleral venous sinus 
= . (canal of Schlemm) 
| 
/ A \ Suspensory ligaments 


Lens 


Lateral rectus 
muscle 


Sclera 


Choroid 


; ———— Cornea 
Retina 


Iris 
] Pupil 
Anterior cavity 
(contains aqueous humor): 
. Posterior chamber 
| J Anterior chamber 
@ 
| r > Suspensory ligaments 


ff [ la Ciliary body: 
' Ciliary process 
Ciliary muscle 


Fovea centralis 


Optic (II) nerve — 


Central retinal 
artery and vein 
Optic disc 
(blind spot) 


Medial rectus 
muscle 


Medial 


The sphere of the eye can be divided into anterior and 
posterior chambers. The wall of the eye is composed of 
three layers: the fibrous tunic, vascular tunic, and neural 

tunic. Within the neural tunic is the retina, with three layers 
of cells and two synaptic layers in between. The center of 
the retina has a small indentation known as the fovea. 


Note that the photoreceptors in the retina (rods and cones) are located behind 
the axons, RGCs, bipolar cells, and retinal blood vessels. A significant amount 
of light is absorbed by these structures before the light reaches the 
photoreceptor cells. However, at the exact center of the retina is a small area 
known as the fovea. At the fovea, the retina lacks the supporting cells and 
blood vessels, and only contains photoreceptors. Therefore, visual acuity, or 
the sharpness of vision, is greatest at the fovea. This is because the fovea is 
where the least amount of incoming light is absorbed by other retinal structures 
(see [link]). As one moves in either direction from this central point of the 
retina, visual acuity drops significantly. In addition, each photoreceptor cell of 
the fovea is connected to a single RGC. Therefore, this RGC does not have to 
integrate inputs from multiple photoreceptors, which reduces the accuracy of 
visual transduction. Toward the edges of the retina, several photoreceptors 


converge on RGCs (through the bipolar cells) up to a ratio of 50 to 1. The 
difference in visual acuity between the fovea and peripheral retina is easily 
evidenced by looking directly at a word in the middle of this paragraph. The 
visual stimulus in the middle of the field of view falls on the fovea and is in the 
sharpest focus. Without moving your eyes off that word, notice that words at 
the beginning or end of the paragraph are not in focus. The images in your 
peripheral vision are focused by the peripheral retina, and have vague, blurry 
edges and words that are not as clearly identified. As a result, a large part of 
the neural function of the eyes is concerned with moving the eyes and head so 
that important visual stimuli are centered on the fovea. 


Light falling on the retina causes chemical changes to pigment molecules in the 
photoreceptors, ultimately leading to a change in the activity of the RGCs. 
Photoreceptor cells have two parts, the inner segment and the outer segment 
([link]). The inner segment contains the nucleus and other common organelles 
of a cell, whereas the outer segment is a specialized region in which 
photoreception takes place. There are two types of photoreceptors—rods and 
cones—which differ in the shape of their outer segment. The rod-shaped outer 
segments of the rod photoreceptor contain a stack of membrane-bound discs 
that contain the photosensitive pigment rhodopsin. The cone-shaped outer 
segments of the cone photoreceptor contain their photosensitive pigments in 
infoldings of the cell membrane. There are three cone photopigments, called 
opsins, which are each sensitive to a particular wavelength of light. The 
wavelength of visible light determines its color. The pigments in human eyes 
are specialized in perceiving three different primary colors: red, green, and 
blue. 

Photoreceptor 


y Pigment 
epithelium 
Melanin granules 


Discs 


Connecting stalks Mitochondria 


Rods 
Golgi apparatus 


Cone 


Nuclei 


Bipolar cell 


Ganglion cell 


(a) 


Choroid 


Pigment epithelium 


Rods and cones 


Bipolar cells 


Ganglion cells 


Optic nerve axons 


(b) 


(a) All photoreceptors have inner segments 
containing the nucleus and other important 
organelles and outer segments with membrane 
arrays containing the photosensitive opsin 
molecules. Rod outer segments are long columnar 
shapes with stacks of membrane-bound discs that 
contain the rhodopsin pigment. Cone outer 
segments are short, tapered shapes with folds of 
membrane in place of the discs in the rods. (b) 


Tissue of the retina shows a dense layer of nuclei 
of the rods and cones. LM x 800. (Micrograph 
provided by the Regents of University of Michigan 
Medical School © 2012) 


At the molecular level, visual stimuli cause changes in the photopigment 
molecule that lead to changes in membrane potential of the photoreceptor cell. 
A single unit of light is called a photon, which is described in physics as a 
packet of energy with properties of both a particle and a wave. The energy of a 
photon is represented by its wavelength, with each wavelength of visible light 
corresponding to a particular color. Visible light is electromagnetic radiation 
with a wavelength between 380 and 720 nm. Wavelengths of electromagnetic 
radiation longer than 720 nm fall into the infrared range, whereas wavelengths 
shorter than 380 nm fall into the ultraviolet range. Light with a wavelength of 
380 nm is blue whereas light with a wavelength of 720 nm is dark red. All 
other colors fall between red and blue at various points along the wavelength 
scale. 


Opsin pigments are actually transmembrane proteins that contain a cofactor 
known as retinal. Retinal is a hydrocarbon molecule related to vitamin A. 
When a photon hits retinal, the long hydrocarbon chain of the molecule is 
biochemically altered. Specifically, photons cause some of the double-bonded 
carbons within the chain to switch from a cis to a trans conformation. This 
process is called photoisomerization. Before interacting with a photon, 
retinal’s flexible double-bonded carbons are in the cis conformation. This 
molecule is referred to as 11-cis-retinal. A photon interacting with the 
molecule causes the flexible double-bonded carbons to change to the trans- 
conformation, forming all-trans-retinal, which has a straight hydrocarbon chain 
((link]). 


The shape change of retinal in the photoreceptors initiates visual transduction 
in the retina. Activation of retinal and the opsin proteins result in activation of 
a G protein. The G protein changes the membrane potential of the 
photoreceptor cell, which then releases less neurotransmitter into the outer 
synaptic layer of the retina. Until the retinal molecule is changed back to the 
11-cis-retinal shape, the opsin cannot respond to light energy, which is called 
bleaching. When a large group of photopigments is bleached, the retina will 


send information as if opposing visual information is being perceived. After a 
bright flash of light, afterimages are usually seen in negative. The 
photoisomerization is reversed by a series of enzymatic changes so that the 
retinal responds to more light energy. 

Retinal Isomers 


Photon 


11-trans-retinal 


11-cis-retinal Fai “eA 
and opsin are 

reassembled 

to form 

rhodopsin 


Regeneration 


Rhodopsin 
molecules 


Rod bar of 


11-cis- tra, 


(a) 11-cis-retinal (b) all-trans-retinal 


The retinal molecule has two isomers, (a) one before a 
photon interacts with it and (b) one that is altered through 
photoisomerization. 


The opsins are sensitive to limited wavelengths of light. Rhodopsin, the 
photopigment in rods, is most sensitive to light at a wavelength of 498 nm. The 
three color opsins have peak sensitivities of 564 nm, 534 nm, and 420 nm 
corresponding roughly to the primary colors of red, green, and blue ({link]). 
The absorbance of rhodopsin in the rods is much more sensitive than in the 


cone opsins; specifically, rods are sensitive to vision in low light conditions, 
and cones are sensitive to brighter conditions. In normal sunlight, rhodopsin 
will be constantly bleached while the cones are active. In a darkened room, 
there is not enough light to activate cone opsins, and vision is entirely 
dependent on rods. Rods are so sensitive to light that a single photon can result 
in an action potential from a rod’s corresponding RGC. 


The three types of cone opsins, being sensitive to different wavelengths of 
light, provide us with color vision. By comparing the activity of the three 
different cones, the brain can extract color information from visual stimuli. For 
example, a bright blue light that has a wavelength of approximately 450 nm 
would activate the “red” cones minimally, the “green” cones marginally, and 
the “blue” cones predominantly. The relative activation of the three different 
cones is calculated by the brain, which perceives the color as blue. However, 
cones cannot react to low-intensity light, and rods do not sense the color of 
light. Therefore, our low-light vision is—in essence—in grayscale. In other 
words, in a dark room, everything appears as a shade of gray. If you think that 
you can see colors in the dark, it is most likely because your brain knows what 
color something is and is relying on that memory. 


Comparison of Color Sensitivity of Photopigments 
420 nm 498nm 534nm 564nm 


Green Red 
Blue cones Rods cones cones 


Normalized absorbance 


400 500 600 700 
Violet Blue Cyan Green Yellow Red 


Wavelength (nm) 


Comparing the peak sensitivity and absorbance 
spectra of the four photopigments suggests that 
they are most sensitive to particular 
wavelengths. 


OR page | 


Watch this video to learn more about a transverse section through the brain 
that depicts the visual pathway from the eye to the occipital cortex. The first 
half of the pathway is the projection from the RGCs through the optic nerve to 
the lateral geniculate nucleus in the thalamus on either side. This first fiber in 
the pathway synapses on a thalamic cell that then projects to the visual cortex 
in the occipital lobe where “seeing,” or visual perception, takes place. This 
video gives an abbreviated overview of the visual system by concentrating on 
the pathway from the eyes to the occipital lobe. The video makes the 
statement (at 0:45) that “specialized cells in the retina called ganglion cells 
convert the light rays into electrical signals.” What aspect of retinal processing 
is simplified by that statement? Explain your answer. 


Sensory Nerves 


Once any sensory cell transduces a stimulus into a nerve impulse, that impulse 
has to travel along axons to reach the CNS. In many of the special senses, the 
axons leaving the sensory receptors have a topographical arrangement, 
meaning that the location of the sensory receptor relates to the location of the 
axon in the nerve. For example, in the retina, axons from RGCs in the fovea 
are located at the center of the optic nerve, where they are surrounded by axons 
from the more peripheral RGCs. 


Spinal Nerves 


Generally, spinal nerves contain afferent axons from sensory receptors in the 
periphery, such as from the skin, mixed with efferent axons travelling to the 
muscles or other effector organs. As the spinal nerve nears the spinal cord, it 


splits into dorsal and ventral roots. The dorsal root contains only the axons of 
sensory neurons, whereas the ventral roots contain only the axons of the motor 
neurons. Some of the branches will synapse with local neurons in the dorsal 
root ganglion, posterior (dorsal) horn, or even the anterior (ventral) horn, at the 
level of the spinal cord where they enter. Other branches will travel a short 
distance up or down the spine to interact with neurons at other levels of the 
spinal cord. A branch may also turn into the posterior (dorsal) column of the 
white matter to connect with the brain. For the sake of convenience, we will 
use the terms ventral and dorsal in reference to structures within the spinal cord 
that are part of these pathways. This will help to underscore the relationships 
between the different components. Typically, spinal nerve systems that connect 
to the brain are contralateral, in that the right side of the body is connected to 
the left side of the brain and the left side of the body to the right side of the 
brain. 


Cranial Nerves 


Cranial nerves convey specific sensory information from the head and neck 
directly to the brain. For sensations below the neck, the right side of the body 
is connected to the left side of the brain and the left side of the body to the right 
side of the brain. Whereas spinal information is contralateral, cranial nerve 
systems are mostly ipsilateral, meaning that a cranial nerve on the right side of 
the head is connected to the right side of the brain. Some cranial nerves contain 
only sensory axons, such as the olfactory, optic, and vestibulocochlear nerves. 
Other cranial nerves contain both sensory and motor axons, including the 
trigeminal, facial, glossopharyngeal, and vagus nerves (however, the vagus 
nerve is not associated with the somatic nervous system). The general senses of 
somatosensation for the face travel through the trigeminal system. 


Chapter Review 


The senses are olfaction (smell), gustation (taste), somatosensation (sensations 
associated with the skin and body), audition (hearing), equilibrium (balance), 
and vision. With the exception of somatosensation, this list represents the 
special senses, or those systems of the body that are associated with specific 
organs such as the tongue or eye. Somatosensation belongs to the general 
senses, which are those sensory structures that are distributed throughout the 


body and in the walls of various organs. The special senses are all primarily 
part of the somatic nervous system in that they are consciously perceived 
through cerebral processes, though some special senses contribute to 
autonomic function. The general senses can be divided into somatosensation, 
which is commonly considered touch, but includes tactile, pressure, vibration, 
temperature, and pain perception. The general senses also include the visceral 
senses, which are separate from the somatic nervous system function in that 
they do not normally rise to the level of conscious perception. 


The cells that transduce sensory stimuli into the electrochemical signals of the 
nervous system are classified on the basis of structural or functional aspects of 
the cells. The structural classifications are either based on the anatomy of the 
cell that is interacting with the stimulus (free nerve endings, encapsulated 
endings, or specialized receptor cell), or where the cell is located relative to the 
stimulus (interoceptor, exteroceptor, proprioceptor). Thirdly, the functional 
classification is based on how the cell transduces the stimulus into a neural 
signal. Chemoreceptors respond to chemical stimuli and are the basis for 
olfaction and gustation. Related to chemoreceptors are osmoreceptors and 
nociceptors for fluid balance and pain reception, respectively. 
Mechanoreceptors respond to mechanical stimuli and are the basis for most 
aspects of somatosensation, as well as being the basis of audition and 
equilibrium in the inner ear. Thermoreceptors are sensitive to temperature 
changes, and photoreceptors are sensitive to light energy. 


The nerves that convey sensory information from the periphery to the CNS are 
either spinal nerves, connected to the spinal cord, or cranial nerves, connected 
to the brain. Spinal nerves have mixed populations of fibers; some are motor 
fibers and some are sensory. The sensory fibers connect to the spinal cord 
through the dorsal root, which is attached to the dorsal root ganglion. Sensory 
information from the body that is conveyed through spinal nerves will project 
to the opposite side of the brain to be processed by the cerebral cortex. The 
cranial nerves can be strictly sensory fibers, such as the olfactory, optic, and 
vestibulocochlear nerves, or mixed sensory and motor nerves, such as the 
trigeminal, facial, glossopharyngeal, and vagus nerves. The cranial nerves are 
connected to the same side of the brain from which the sensory information 
originates. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to learn about Dr. Danielle Reed of the Monell Chemical 
Senses Center in Philadelphia, PA, who became interested in science at an 
early age because of her sensory experiences. She recognized that her 
sense of taste was unique compared with other people she knew. Now, she 
studies the genetic differences between people and their sensitivities to 
taste stimuli. In the video, there is a brief image of a person sticking out 
their tongue, which has been covered with a colored dye. This is how Dr. 
Reed is able to visualize and count papillae on the surface of the tongue. 
People fall into two large groups known as “tasters” and “non-tasters” on 
the basis of the density of papillae on their tongue, which also indicates 
the number of taste buds. Non-tasters can taste food, but they are not as 
sensitive to certain tastes, such as bitterness. Dr. Reed discovered that she 
is anon-taster, which explains why she perceived bitterness differently 
than other people she knew. Are you very sensitive to tastes? Can you see 
any similarities among the members of your family? 


Solution: 


Answers will vary, but a typical answer might be: I can eat most anything 
(except mushrooms! ), so I don’t think that I’m that sensitive to tastes. My 
whole family likes eating a variety of foods, so it seems that we all have 
the same level of sensitivity. 


Exercise: 
Problem: 
[link] The basilar membrane is the thin membrane that extends from the 
central core of the cochlea to the edge. What is anchored to this 


membrane so that they can be activated by movement of the fluids within 
the cochlea? 


Solution: 


[link] The hair cells are located in the organ of Corti, which is located on 
the basilar membrane. The stereocilia of those cells would normally be 


attached to the tectorial membrane (though they are detached in the 
micrograph because of processing of the tissue). 


Exercise: 


Problem: 


Watch this video to learn more about how the structures of the ear convert 
sound waves into a neural signal by moving the “hairs,” or stereocilia, of 
the cochlear duct. Specific locations along the length of the duct encode 
specific frequencies, or pitches. The brain interprets the meaning of the 
sounds we hear as music, speech, noise, etc. Which ear structures are 
responsible for the amplification and transfer of sound from the external 
ear to the inner ear? 


Solution: 


The small bones in the middle ear, the ossicles, amplify and transfer 
sound between the tympanic membrane of the external ear and the oval 
window of the inner ear. 


Exercise: 


Problem: 


Watch this animation to learn more about the inner ear and to see the 
cochlea unroll, with the base at the back of the image and the apex at the 
front. Specific wavelengths of sound cause specific regions of the basilar 
membrane to vibrate, much like the keys of a piano produce sound at 
different frequencies. Based on the animation, where do frequencies— 
from high to low pitches—cause activity in the hair cells within the 
cochlear duct? 


Solution: 


High frequencies activate hair cells toward the base of the cochlea, and 
low frequencies activate hair cells toward the apex of the cochlea. 


Exercise: 


Problem: 


Watch this video to learn more about a transverse section through the 
brain that depicts the visual pathway from the eye to the occipital cortex. 
The first half of the pathway is the projection from the RGCs through the 
optic nerve to the lateral geniculate nucleus in the thalamus on either side. 
This first fiber in the pathway synapses on a thalamic cell that then 
projects to the visual cortex in the occipital lobe where “seeing,” or visual 
perception, takes place. This video gives an abbreviated overview of the 
visual system by concentrating on the pathway from the eyes to the 
occipital lobe. The video makes the statement (at 0:45) that “specialized 
cells in the retina called ganglion cells convert the light rays into electrical 
signals.” What aspect of retinal processing is simplified by that statement? 
Explain your answer. 


Solution: 


Photoreceptors convert light energy, or photons, into an electrochemical 
signal. The retina contains bipolar cells and the RGCs that finally convert 
it into action potentials that are sent from the retina to the CNS. It is 
important to recognize when popular media and online sources 
oversimplify complex physiological processes so that misunderstandings 
are not generated. This video was created by a medical device 
manufacturer who might be trying to highlight other aspects of the visual 
system than retinal processing. The statement they make is not incorrect, 
it just bundles together several steps, which makes it sound like RGCs are 
the transducers, rather than photoreceptors. 


Review Questions 


Exercise: 


Problem: 
What type of receptor cell is responsible for transducing pain stimuli? 


a. mechanoreceptor 
b. nociceptor 
c. osmoreceptor 


d. photoreceptor 


Solution: 


B 


Exercise: 


Problem: Which of these cranial nerves is part of the gustatory system? 


a. olfactory 
b. trochlear 
c. trigeminal 
d. facial 


Solution: 
D 


Exercise: 


Problem:Which submodality of taste is sensitive to the pH of saliva? 


a. umMami 
b. sour 

c. bitter 
d. sweet 


Solution: 


B 


Exercise: 


Problem: Axons from which neuron in the retina make up the optic nerve? 


a. amacrine cells 
b. photoreceptors 


c. bipolar cells 
d. retinal ganglion cells 


Solution: 


D 
Exercise: 


Problem: 


What type of receptor cell is involved in the sensations of sound and 
balance? 


a. photoreceptor 

b. chemoreceptor 

c. mechanoreceptor 
d. nociceptor 


Solution: 


C 


Critical Thinking Questions 


Exercise: 
Problem: 


The sweetener known as stevia can replace glucose in food. What does 
the molecular similarity of stevia to glucose mean for the gustatory sense? 


Solution: 


The stevia molecule is similar to glucose such that it will bind to the 
glucose receptor in sweet-sensitive taste buds. However, it is not a 
substrate for the ATP-generating metabolism within cells. 


Exercise: 


Problem: 


Why does the blind spot from the optic disc in either eye not result in a 
blind spot in the visual field? 


Solution: 


The visual field for each eye is projected onto the retina as light is focused 
by the lens. The visual information from the right visual field falls on the 
left side of the retina and vice versa. The optic disc in the right eye is on 
the medial side of the fovea, which would be the left side of the retina. 
However, the optic disc in the left eye would be on the right side of that 
fovea, so the right visual field falls on the side of the retina in the left field 
where there is no blind spot. 


Glossary 


alkaloid 
substance, usually from a plant source, that is chemically basic with 
respect to pH and will stimulate bitter receptors 


amacrine cell 
type of cell in the retina that connects to the bipolar cells near the outer 
synaptic layer and provides the basis for early image processing within 
the retina 


ampulla 
in the ear, the structure at the base of a semicircular canal that contains the 
hair cells and cupula for transduction of rotational movement of the head 


anosmia 
loss of the sense of smell; usually the result of physical disruption of the 
first cranial nerve 


aqueous humor 
watery fluid that fills the anterior chamber containing the cornea, iris, 
ciliary body, and lens of the eye 


audition 
sense of hearing 


auricle 
fleshy external structure of the ear 


basilar membrane 
in the ear, the floor of the cochlear duct on which the organ of Corti sits 


bipolar cell 
cell type in the retina that connects the photoreceptors to the RGCs 


capsaicin 
molecule that activates nociceptors by interacting with a temperature- 
sensitive ion channel and is the basis for “hot” sensations in spicy food 


chemoreceptor 
sensory receptor cell that is sensitive to chemical stimuli, such as in taste, 
smell, or pain 


choroid 
highly vascular tissue in the wall of the eye that supplies the outer retina 
with blood 


ciliary body 
smooth muscle structure on the interior surface of the iris that controls the 
shape of the lens through the zonule fibers 


cochlea 
auditory portion of the inner ear containing structures to transduce sound 
stimuli 


cochlear duct 
space within the auditory portion of the inner ear that contains the organ 
of Corti and is adjacent to the scala tympani and scala vestibuli on either 
side 


cone photoreceptor 
one of the two types of retinal receptor cell that is specialized for color 
vision through the use of three photopigments distributed through three 


separate populations of cells 


contralateral 
word meaning “on the opposite side,” as in axons that cross the midline in 
a fiber tract 


cornea 
fibrous covering of the anterior region of the eye that is transparent so that 
light can pass through it 


cupula 
specialized structure within the base of a semicircular canal that bends the 
stereocilia of hair cells when the head rotates by way of the relative 
movement of the enclosed fluid 


encapsulated ending 
configuration of a sensory receptor neuron with dendrites surrounded by 
specialized structures to aid in transduction of a particular type of 
sensation, such as the lamellated corpuscles in the deep dermis and 
subcutaneous tissue 


equilibrium 
sense of balance that includes sensations of position and movement of the 
head 


external ear 
structures on the lateral surface of the head, including the auricle and the 
ear canal back to the tympanic membrane 


exteroceptor 
sensory receptor that is positioned to interpret stimuli from the external 
environment, such as photoreceptors in the eye or somatosensory 
receptors in the skin 


extraocular muscle 
one of six muscles originating out of the bones of the orbit and inserting 
into the surface of the eye which are responsible for moving the eye 


fibrous tunic 


outer layer of the eye primarily composed of connective tissue known as 
the sclera and cornea 


fovea 
exact center of the retina at which visual stimuli are focused for maximal 
acuity, where the retina is thinnest, at which there is nothing but 
photoreceptors 


free nerve ending 
configuration of a sensory receptor neuron with dendrites in the 
connective tissue of the organ, such as in the dermis of the skin, that are 
most often sensitive to chemical, thermal, and mechanical stimuli 


general sense 
any sensory system that is distributed throughout the body and 
incorporated into organs of multiple other systems, such as the walls of 
the digestive organs or the skin 


gustation 
sense of taste 


gustatory receptor cells 
sensory cells in the taste bud that transduce the chemical stimuli of 
gustation 


hair cells 
mechanoreceptor cells found in the inner ear that transduce stimuli for the 
senses of hearing and balance 


incus 
(also, anvil) ossicle of the middle ear that connects the malleus to the 
stapes 


inferior oblique 
extraocular muscle responsible for lateral rotation of the eye 


inferior rectus 
extraocular muscle responsible for looking down 


inner ear 


structure within the temporal bone that contains the sensory apparati of 
hearing and balance 


inner segment 
in the eye, the section of a photoreceptor that contains the nucleus and 
other major organelles for normal cellular functions 


inner synaptic layer 
layer in the retina where bipolar cells connect to RGCs 


interoceptor 
sensory receptor that is positioned to interpret stimuli from internal 
organs, such as stretch receptors in the wall of blood vessels 


ipsilateral 
word meaning on the same side, as in axons that do not cross the midline 
in a fiber tract 


iris 
colored portion of the anterior eye that surrounds the pupil 
kinesthesia 


sense of body movement based on sensations in skeletal muscles, tendons, 
joints, and the skin 


lacrimal duct 
duct in the medial corner of the orbit that drains tears into the nasal cavity 


lacrimal gland 
gland lateral to the orbit that produces tears to wash across the surface of 
the eye 


lateral rectus 
extraocular muscle responsible for abduction of the eye 


lens 
component of the eye that focuses light on the retina 


levator palpebrae superioris 


muscle that causes elevation of the upper eyelid, controlled by fibers in 
the oculomotor nerve 


macula 
enlargement at the base of a semicircular canal at which transduction of 
equilibrium stimuli takes place within the ampulla 


malleus 
(also, hammer) ossicle that is directly attached to the tympanic membrane 


mechanoreceptor 
receptor cell that transduces mechanical stimuli into an electrochemical 
signal 


medial rectus 
extraocular muscle responsible for adduction of the eye 


middle ear 
space within the temporal bone between the ear canal and bony labyrinth 
where the ossicles amplify sound waves from the tympanic membrane to 
the oval window 


neural tunic 
layer of the eye that contains nervous tissue, namely the retina 


nociceptor 
receptor cell that senses pain stimuli 


odorant molecules 
volatile chemicals that bind to receptor proteins in olfactory neurons to 
stimulate the sense of smell 


olfaction 
sense of smell 


olfactory bulb 
central target of the first cranial nerve; located on the ventral surface of 
the frontal lobe in the cerebrum 


olfactory epithelium 


region of the nasal epithelium where olfactory neurons are located 


olfactory sensory neuron 
receptor cell of the olfactory system, sensitive to the chemical stimuli of 
smell, the axons of which compose the first cranial nerve 


opsin 
protein that contains the photosensitive cofactor retinal for 
phototransduction 


optic disc 
spot on the retina at which RGC axons leave the eye and blood vessels of 
the inner retina pass 


optic nerve 
second cranial nerve, which is responsible visual sensation 


organ of Corti 
structure in the cochlea in which hair cells transduce movements from 
sound waves into electrochemical signals 


osmoreceptor 
receptor cell that senses differences in the concentrations of bodily fluids 
on the basis of osmotic pressure 


ossicles 
three small bones in the middle ear 


otolith 
layer of calcium carbonate crystals located on top of the otolithic 
membrane 


otolithic membrane 
gelatinous substance in the utricle and saccule of the inner ear that 
contains calcium carbonate crystals and into which the stereocilia of hair 
cells are embedded 


outer segment 
in the eye, the section of a photoreceptor that contains opsin molecules 
that transduce light stimuli 


outer synaptic layer 
layer in the retina at which photoreceptors connect to bipolar cells 


oval window 
membrane at the base of the cochlea where the stapes attaches, marking 
the beginning of the scala vestibuli 


palpebral conjunctiva 
membrane attached to the inner surface of the eyelids that covers the 
anterior surface of the cornea 


papilla 
for gustation, a bump-like projection on the surface of the tongue that 
contains taste buds 


photoisomerization 
chemical change in the retinal molecule that alters the bonding so that it 
switches from the 11-cis-retinal isomer to the all-trans-retinal isomer 


photon 
individual “packet” of light 


photoreceptor 
receptor cell specialized to respond to light stimuli 


proprioception 
sense of position and movement of the body 


proprioceptor 
receptor cell that senses changes in the position and kinesthetic aspects of 
the body 


pupil 
open hole at the center of the iris that light passes through into the eye 


receptor cell 
cell that transduces environmental stimuli into neural signals 


retina 
nervous tissue of the eye at which phototransduction takes place 


retinal 
cofactor in an opsin molecule that undergoes a biochemical change when 
struck by a photon (pronounced with a stress on the last syllable) 


retinal ganglion cell (RGC) 
neuron of the retina that projects along the second cranial nerve 


rhodopsin 
photopigment molecule found in the rod photoreceptors 


rod photoreceptor 
one of the two types of retinal receptor cell that is specialized for low- 
light vision 


round window 
membrane that marks the end of the scala tympani 


saccule 
structure of the inner ear responsible for transducing linear acceleration in 
the vertical plane 


scala tympani 
portion of the cochlea that extends from the apex to the round window 


scala vestibuli 
portion of the cochlea that extends from the oval window to the apex 


sclera 
white of the eye 


semicircular canals 
structures within the inner ear responsible for transducing rotational 
movement information 


sensory modality 
a particular system for interpreting and perceiving environmental stimuli 
by the nervous system 


somatosensation 
general sense associated with modalities lumped together as touch 


special sense 
any sensory system associated with a specific organ structure, namely 
smell, taste, sight, hearing, and balance 


spiral ganglion 
location of neuronal cell bodies that transmit auditory information along 
the eighth cranial nerve 


stapes 
(also, stirrup) ossicle of the middle ear that is attached to the inner ear 


stereocilia 
array of apical membrane extensions in a hair cell that transduce 
movements when they are bent 


submodality 
specific sense within a broader major sense such as sweet as a part of the 
sense of taste, or color as a part of vision 


superior oblique 
extraocular muscle responsible for medial rotation of the eye 


superior rectus 
extraocular muscle responsible for looking up 


taste buds 
structures within a papilla on the tongue that contain gustatory receptor 
cells 


tectorial membrane 
component of the organ of Corti that lays over the hair cells, into which 
the stereocilia are embedded 


thermoreceptor 
sensory receptor specialized for temperature stimuli 


topographical 
relating to positional information 


transduction 


process of changing an environmental stimulus into the electrochemical 
signals of the nervous system 


trochlea 
cartilaginous structure that acts like a pulley for the superior oblique 
muscle 


tympanic membrane 
ear drum 


umami 
taste submodality for sensitivity to the concentration of amino acids; also 
called the savory sense 


utricle 
structure of the inner ear responsible for transducing linear acceleration in 
the horizontal plane 


vascular tunic 
middle layer of the eye primarily composed of connective tissue with a 
rich blood supply 


vestibular ganglion 
location of neuronal cell bodies that transmit equilibrium information 
along the eighth cranial nerve 


vestibule 
in the ear, the portion of the inner ear responsible for the sense of 
equilibrium 


visceral sense 
sense associated with the internal organs 


vision 
special sense of sight based on transduction of light stimuli 
visual acuity 


property of vision related to the sharpness of focus, which varies in 
relation to retinal position 


vitreous humor 
viscous fluid that fills the posterior chamber of the eye 


zonule fibers 
fibrous connections between the ciliary body and the lens 


Divisions of the ANS 
By the end of this section, you will be able to: 


e Name the components that generate the sympathetic and 
parasympathetic responses of the autonomic nervous system 

e Explain the differences in output connections within the two divisions 
of the autonomic nervous system 

e Describe the signaling molecules and receptor proteins involved in 
communication within the two divisions of the autonomic nervous 
system 


The nervous system can be divided into two functional parts: the somatic 
nervous system and the autonomic nervous system. The major differences 
between the two systems are evident in the responses that each produces. 
The somatic nervous system causes contraction of skeletal muscles. The 
autonomic nervous system controls cardiac and smooth muscle, as well as 
glandular tissue. The somatic nervous system is associated with voluntary 
responses (though many can happen without conscious awareness, like 
breathing), and the autonomic nervous system is associated with 
involuntary responses, such as those related to homeostasis. 


The autonomic nervous system regulates many of the internal organs 
through a balance of two aspects, or divisions. In addition to the endocrine 
system, the autonomic nervous system is instrumental in homeostatic 
mechanisms in the body. The two divisions of the autonomic nervous 
system are the sympathetic division and the parasympathetic division. 
The sympathetic system is associated with the fight-or-flight response, and 
parasympathetic activity is referred to by the epithet of rest and digest. 
Homeostasis is the balance between the two systems. At each target 
effector, dual innervation determines activity. For example, the heart 
receives connections from both the sympathetic and parasympathetic 
divisions. One causes heart rate to increase, whereas the other causes heart 
rate to decrease. 


Note: 


— 
wees Openstax COLLEGE 


ie Batt 


Watch this video to learn more about adrenaline and the fight-or-flight 
response. When someone is said to have a rush of adrenaline, the image of 
bungee jumpers or skydivers usually comes to mind. But adrenaline, also 
known as epinephrine, is an important chemical in coordinating the body’s 
fight-or-flight response. In this video, you look inside the physiology of the 
fight-or-flight response, as envisioned for a firefighter. His body’s reaction 
is the result of the sympathetic division of the autonomic nervous system 
causing system-wide changes as it prepares for extreme responses. What 
two changes does adrenaline bring about to help the skeletal muscle 
response? 


Sympathetic Division of the Autonomic Nervous System 


To respond to a threat—to fight or to run away—the sympathetic system 
causes divergent effects as many different effector organs are activated 
together for a common purpose. More oxygen needs to be inhaled and 
delivered to skeletal muscle. The respiratory, cardiovascular, and 
musculoskeletal systems are all activated together. Additionally, sweating 
keeps the excess heat that comes from muscle contraction from causing the 
body to overheat. The digestive system shuts down so that blood is not 
absorbing nutrients when it should be delivering oxygen to skeletal 
muscles. To coordinate all these responses, the connections in the 
sympathetic system diverge from a limited region of the central nervous 
system (CNS) to a wide array of ganglia that project to the many effector 
organs simultaneously. The complex set of structures that compose the 
output of the sympathetic system make it possible for these disparate 
effectors to come together in a coordinated, systemic change. 


The sympathetic division of the autonomic nervous system influences the 
various organ systems of the body through connections emerging from the 
thoracic and upper lumbar spinal cord. It is referred to as the 
thoracolumbar system to reflect this anatomical basis. A central neuron 
in the lateral horn of any of these spinal regions projects to ganglia adjacent 
to the vertebral column through the ventral spinal roots. The majority of 
ganglia of the sympathetic system belong to a network of sympathetic 
chain ganglia that runs alongside the vertebral column. The ganglia appear 
as a series of clusters of neurons linked by axonal bridges. There are 
typically 23 ganglia in the chain on either side of the spinal column. Three 
correspond to the cervical region, 12 are in the thoracic region, four are in 
the lumbar region, and four correspond to the sacral region. The cervical 
and sacral levels are not connected to the spinal cord directly through the 
spinal roots, but through ascending or descending connections through the 
bridges within the chain. 


A diagram that shows the connections of the sympathetic system is 
somewhat like a circuit diagram that shows the electrical connections 
between different receptacles and devices. In [link], the “circuits” of the 
sympathetic system are intentionally simplified. 

Connections of Sympathetic Division of the Autonomic Nervous System 


Region of 
spinal cord 


Left chain 
ganglia 


Medulla 


ee 
‘ # Ze 
Right chain ‘ « .o 
ganglia gee gi 
WX 7° 1-7 cet 
2 ex\0 
¢} ey? 


mM 


Spinal cord a 


Coccygeal ganglia 
fused together 
(ganglion impar) 


} 


Associated nerves 
and prevertebral ganglia 


eee eee cress 
oc 


Superior 
mesenteric 
ganglion 


i 


Inferior 


ganglion 


3 Celiac 
ganglion 


Cs) . 
% 
. 
“ x 
mesenteric N14 “S 


Target organs (effectors) 


Lacrinal gland 


-“‘G@eepEp Mucous membrane- 
Siete nose and palate 


Submaxillary gland 
~~~ Gaex Sublingual gland 


= Mucous membrane- 
mouth 


Parotid gland 


Heart 


Larynx 
Trachea 


Bronchi 
Esophagus 


Stomach 


Abdominal 
blood vessels 


Large 
intestine 


Rectum 


Kidney 


Bladder 


Gonads 


External genitalia 


Neurons from the lateral horn of the spinal cord 
(preganglionic nerve fibers - solid lines)) project to the 
chain ganglia on either side of the vertebral column or to 


collateral (prevertebral) ganglia that are anterior to the 
vertebral column in the abdominal cavity. Axons from 
these ganglionic neurons (postganglionic nerve fibers - 
dotted lines) then project to target effectors throughout the 
body. 


To continue with the analogy of the circuit diagram, there are three different 
types of “junctions” that operate within the sympathetic system ([link]). The 
first type is most direct: the sympathetic nerve projects to the chain 
ganglion at the same level as the target effector (the organ, tissue, or gland 
to be innervated). An example of this type is spinal nerve T1 that synapses 
with the T1 chain ganglion to innervate the trachea. The fibers of this 
branch are called white rami communicantes (singular = ramus 
communicans); they are myelinated and therefore referred to as white (see 
[link]a). The axon from the central neuron (the preganglionic fiber shown 
as a Solid line) synapses with the ganglionic neuron (with the 
postganglionic fiber shown as a dashed line). This neuron then projects to a 
target effector—in this case, the trachea—via gray rami communicantes, 
which are unmyelinated axons. 


In some cases, the target effectors are located superior or inferior to the 
spinal segment at which the preganglionic fiber emerges. With respect to 
the “wiring” involved, the synapse with the ganglionic neuron occurs at 
chain ganglia superior or inferior to the location of the central neuron. An 
example of this is spinal nerve T1 that innervates the eye. The spinal nerve 
tracks up through the chain until it reaches the superior cervical ganglion, 
where it synapses with the postganglionic neuron (see [link |b). The cervical 
ganglia are referred to as paravertebral ganglia, given their location 
adjacent to prevertebral ganglia in the sympathetic chain. 


Not all axons from the central neurons terminate in the chain ganglia. 
Additional branches from the ventral nerve root continue through the chain 
and on to one of the collateral ganglia as the greater splanchnic nerve or 
lesser splanchnic nerve. For example, the greater splanchnic nerve at the 
level of TS synapses with a collateral ganglion outside the chain before 


making the connection to the postganglionic nerves that innervate the 
stomach (see [link]c). 


Collateral ganglia, also called prevertebral ganglia, are situated anterior 
to the vertebral column and receive inputs from splanchnic nerves as well 
as central sympathetic neurons. They are associated with controlling organs 
in the abdominal cavity, and are also considered part of the enteric nervous 
system. The three collateral ganglia are the celiac ganglion, the superior 
mesenteric ganglion, and the inferior mesenteric ganglion (see [link]). 
The word celiac is derived from the Latin word “coelom,” which refers to a 
body cavity (in this case, the abdominal cavity), and the word mesenteric 
refers to the digestive system. 

Sympathetic Connections and Chain Ganglia 


(a) A central neuron synapses 
with a ganglion at the same z 
level within the chain ganglia. a ) Sympathetic 
chain ganglion 


; White ramus 
Spinal | communicans 
Spinal cord —_ nerve ql 
---= To target effector 


Dorsal root 
ganglion 


\ To target effector 


| BZ Gray ramus 
(b) A central neuron N 4 ; : communicans 
synapses within a : } AN 
more superior or inferior Ww 


Spinal 
ganglion in the chain. Seb 


Spinal cord nerve 


(c) Acentral neuron projects 
through the white ramus but 
does not synapse in a chain 
ganglion. Instead, it continues 
through one of the splanchnic 
nerves to synapse within a 
prevertebral ganglion. 


Sympathetic 
chain ganglion 


2 No synapse in 
Spinal . spinal ganglion 
inal cor ner ; 
apineheere ne Prevertebral 


: ~Sx. ganglion 


/~ 
To target effector 


Splanchnic nerve 


Axon of central neuron 
---- Axon of ganglionic neuron 
e Central neuron body 
©  Ganglionic neuron body 
—© Synapse 


The axon from a central sympathetic neuron in the spinal 

cord can project to the periphery in a number of different 

ways. (a) The fiber can project out to the ganglion at the 

same level and synapse on a ganglionic neuron. (b) A 

branch can project to more superior or inferior ganglion in 
the chain. (c) A branch can project through the white ramus 
communicans, but not terminate on a ganglionic neuron in 
the chain. Instead, it projects through one of the splanchnic 


nerves to a collateral ganglion or the adrenal medulla (not 
pictured). 


An axon from the central neuron that projects to a sympathetic ganglion is 
referred to as a preganglionic fiber or neuron, and represents the output 
from the CNS to the ganglion. Because the sympathetic ganglia are adjacent 
to the vertebral column, preganglionic sympathetic fibers are relatively 
short, and they are myelinated. A postganglionic fiber—the axon from a 
ganglionic neuron that projects to the target effector—represents the output 
of a ganglion that directly influences the organ. Compared with the 
preganglionic fibers, postganglionic sympathetic fibers are long because of 
the relatively greater distance from the ganglion to the target effector. These 
fibers are unmyelinated. (Note that the term “postganglionic neuron” may 
be used to describe the projection from a ganglion to the target. The 
problem with that usage is that the cell body is in the ganglion, and only the 
fiber is postganglionic. Typically, the term neuron applies to the entire cell.) 


One type of preganglionic sympathetic fiber does not terminate in a 
ganglion. These are the axons from central sympathetic neurons that project 
to the adrenal medulla, the interior portion of the adrenal gland. These 
axons are still referred to as preganglionic fibers, but the target is not a 
ganglion. The adrenal medulla releases signaling molecules into the 
bloodstream, rather than using axons to communicate with target structures. 
The cells in the adrenal medulla that are contacted by the preganglionic 
fibers are called chromaffin cells. These cells are neurosecretory cells that 
develop from the neural crest along with the sympathetic ganglia, 
reinforcing the idea that the gland is, functionally, a sympathetic ganglion. 


The projections of the sympathetic division of the autonomic nervous 
system diverge widely, resulting in a broad influence of the system 
throughout the body. As a response to a threat, the sympathetic system 
would increase heart rate and breathing rate and cause blood flow to the 
skeletal muscle to increase and blood flow to the digestive system to 
decrease. Sweat gland secretion should also increase as part of an integrated 
response. All of those physiological changes are going to be required to 
occur together to run away from the hunting lioness, or the modern 
equivalent. This divergence is seen in the branching patterns of 


preganglionic sympathetic neurons—a single preganglionic sympathetic 
neuron may have 10—20 targets. An axon that leaves a central neuron of the 
lateral horn in the thoracolumbar spinal cord will pass through the white 
ramus communicans and enter the sympathetic chain, where it will branch 
toward a variety of targets. At the level of the spinal cord at which the 
preganglionic sympathetic fiber exits the spinal cord, a branch will synapse 
on a neuron in the adjacent chain ganglion. Some branches will extend up 
or down to a different level of the chain ganglia. Other branches will pass 
through the chain ganglia and project through one of the splanchnic nerves 
to a collateral ganglion. Finally, some branches may project through the 
splanchnic nerves to the adrenal medulla. All of these branches mean that 
one preganglionic neuron can influence different regions of the sympathetic 
system very broadly, by acting on widely distributed organs. 


Parasympathetic Division of the Autonomic Nervous System 


The parasympathetic division of the autonomic nervous system is named 
because its central neurons are located on either side of the thoracolumbar 
region of the spinal cord (para- = “beside” or “near”). The parasympathetic 
system can also be referred to as the craniosacral system (or outflow) 
because the preganglionic neurons are located in nuclei of the brain stem 
and the lateral horn of the sacral spinal cord. 


The connections, or “circuits,” of the parasympathetic division are similar 
to the general layout of the sympathetic division with a few specific 
differences ([{link]). The preganglionic fibers from the cranial region travel 
in cranial nerves, whereas preganglionic fibers from the sacral region travel 
in spinal nerves. The targets of these fibers are terminal ganglia, which are 
located near—or even within—the target effector. These ganglia are often 
referred to as intramural ganglia when they are found within the walls of 
the target organ. The postganglionic fiber projects from the terminal ganglia 
a short distance to the target effector, or to the specific target tissue within 
the organ. Comparing the relative lengths of axons in the parasympathetic 
system, the preganglionic fibers are long and the postganglionic fibers are 
short because the ganglia are close to—and sometimes within—the target 
effectors. 


The cranial component of the parasympathetic system is based in particular 
nuclei of the brain stem. In the midbrain, the Edinger—Westphal nucleus is 
part of the oculomotor complex, and axons from those neurons travel with 
the fibers in the oculomotor nerve (cranial nerve III) that innervate the 
extraocular muscles. The preganglionic parasympathetic fibers within 
cranial nerve III terminate in the ciliary ganglion, which is located in the 
posterior orbit. The postganglionic parasympathetic fibers then project to 
the smooth muscle of the iris to control pupillary size. In the upper medulla, 
the salivatory nuclei contain neurons with axons that project through the 
facial and glossopharyngeal nerves to ganglia that control salivary glands. 
Tear production is influenced by parasympathetic fibers in the facial nerve, 
which activate a ganglion, and ultimately the lacrimal (tear) gland. Neurons 
in the dorsal nucleus of the vagus nerve and the nucleus ambiguus 
project through the vagus nerve (cranial nerve X) to the terminal ganglia of 
the thoracic and abdominal cavities. Parasympathetic preganglionic fibers 
primarily influence the heart, bronchi, and esophagus in the thoracic cavity 
and the stomach, liver, pancreas, gall bladder, and small intestine of the 
abdominal cavity. The postganglionic fibers from the ganglia activated by 
the vagus nerve are often incorporated into the structure of the organ, such 
as the mesenteric plexus of the digestive tract organs and the intramural 
ganglia. 

Connections of Parasympathetic Division of the Autonomic Nervous 
System 


Eddinger—Westpha 
nucleus 


Super salivatory 
nucleus 


Inferior salivatory 
nucleus 


Dorsal nucleus 
of the vagus and 
nucleus ambiguus 


Oe ee ee es 


Spinal cord 


Region of Associated nerves Target organs (effectors) 
spinal cord and terminal ganglia 


Ciliary ganglion a é , 
A __\ Cranial nerve Ill eS #) 
SS Pterygopalatine 

| 


Cranial nerve VII ganglion G2) =) 
\ 


—} Submandibular 
ganglion 


S5 


0 \ 
AJ Coccygeal v 
ganglia 


fused together DreoyG@ 
(ganglion impar) 
a 


Parasympathetic fibers 


—— Sympathetic fibers 


Eye 


Lacrinal gland 


Mucous membrane 
(nose and palate) 


Submaxillary gland 


Sublingual gland 


Mucous membrane 
(mouth) 


Parotid gland 


Heart 


Larynx 
Trachea 
Bronchi 


Esophagus 


Stomach 


Abdominal blood 
vessels 


Liver and bile duct 


Pancreas 


Adrenal gland 


Small intestine 


Large intestine 


Rectum 


Kidney 
Bladder 


Gonads 


External genitalia 


Neurons from brain-stem nuclei, or from the lateral horn of 


the sacral spinal cord, project to terminal ganglia near or 
within the various organs of the body. Axons from these 
ganglionic neurons then project the short distance to those 
target effectors. 


Chemical Signaling in the Autonomic Nervous System 


Where an autonomic neuron connects with a target, there is a synapse. The 
electrical signal of the action potential causes the release of a signaling 
molecule, which will bind to receptor proteins on the target cell. Synapses 
of the autonomic system are classified as either cholinergic, meaning that 
acetylcholine (ACh) is released, or adrenergic, meaning that 
norepinephrine is released. The terms cholinergic and adrenergic refer not 
only to the signaling molecule that is released but also to the class of 
receptors that each binds. 


The cholinergic system includes two classes of receptor: the nicotinic 
receptor and the muscarinic receptor. Both receptor types bind to ACh 
and cause changes in the target cell. The nicotinic receptor is a ligand- 
gated cation channel and the muscarinic receptor is a G protein—coupled 
receptor. The receptors are named for, and differentiated by, other 
molecules that bind to them. Whereas nicotine will bind to the nicotinic 
receptor, and muscarine will bind to the muscarinic receptor, there is no 
cross-reactivity between the receptors. The situation is similar to locks and 
keys. Imagine two locks—one for a classroom and the other for an office— 
that are opened by two separate keys. The classroom key will not open the 
office door and the office key will not open the classroom door. This is 
similar to the specificity of nicotine and muscarine for their receptors. 
However, a master key can open multiple locks, such as a master key for 
the Biology Department that opens both the classroom and the office doors. 
This is similar to ACh that binds to both types of receptors. The molecules 
that define these receptors are not crucial—they are simply tools for 
researchers to use in the laboratory. These molecules are exogenous, 
meaning that they are made outside of the human body, so a researcher can 


use them without any confounding endogenous results (results caused by 
the molecules produced in the body). 


The adrenergic system also has two types of receptors, named the alpha 
(a)-adrenergic receptor and beta (f)-adrenergic receptor. Unlike 
cholinergic receptors, these receptor types are not classified by which drugs 
can bind to them. All of them are G protein-coupled receptors. There are 
three types of a-adrenergic receptors, termed a1, Q, and a3, and there are 
two types of f-adrenergic receptors, termed 8, and B5. An additional aspect 
of the adrenergic system is that there is a second signaling molecule called 
epinephrine. The chemical difference between norepinephrine and 
epinephrine is the addition of a methyl group (CH3) in epinephrine. The 
prefix “nor-” actually refers to this chemical difference, in which a methyl 
group is missing. 


The term adrenergic should remind you of the word adrenaline, which is 
associated with the fight-or-flight response described at the beginning of the 
chapter. Adrenaline and epinephrine are two names for the same molecule. 
The adrenal gland (in Latin, ad- = “on top of”; renal = “kidney”) secretes 
adrenaline. The ending “-ine” refers to the chemical being derived, or 
extracted, from the adrenal gland. A similar construction from Greek 
instead of Latin results in the word epinephrine (epi- = “above”; nephr- = 
“kidney”). In scientific usage, epinephrine is preferred in the United States, 
whereas adrenaline is preferred in Great Britain, because “adrenalin” was 
once a registered, proprietary drug name in the United States. Though the 
drug is no longer sold, the convention of referring to this molecule by the 
two different names persists. Similarly, norepinephrine and noradrenaline 
are two names for the same molecule. 


Having understood the cholinergic and adrenergic systems, their role in the 
autonomic system is relatively simple to understand. All preganglionic 
fibers, both sympathetic and parasympathetic, release ACh. All ganglionic 
neurons—the targets of these preganglionic fibers—have nicotinic receptors 
in their cell membranes. The nicotinic receptor is a ligand-gated cation 
channel that results in depolarization of the postsynaptic membrane. The 
postganglionic parasympathetic fibers also release ACh, but the receptors 
on their targets are muscarinic receptors, which are G protein—coupled 


receptors and do not exclusively cause depolarization of the postsynaptic 
membrane. Postganglionic sympathetic fibers release norepinephrine, 
except for fibers that project to sweat glands and to blood vessels associated 
with skeletal muscles, which release ACh ((link]). 


Autonomic System Signaling Molecules 


Sympathetic Parasympathetic 
hs Acetylcholine > Acetylcholine > 
Preganglionic ete tay een 
nicotinic receptor nicotinic receptor 
Norepinephrine — a- or 
B-adrenergic receptors 
Acetylcholine > 
secon Acetylcholine > 
a muscarinic receptor a 
Postganglionic muscarinic 
(associated with sweat 
receptor 


glands and the blood 
vessels associated with 
skeletal muscles only 


Signaling molecules can belong to two broad groups. Neurotransmitters are 
released at synapses, whereas hormones are released into the bloodstream. 
These are simplistic definitions, but they can help to clarify this point. 
Acetylcholine can be considered a neurotransmitter because it is released by 
axons at synapses. The adrenergic system, however, presents a challenge. 
Postganglionic sympathetic fibers release norepinephrine, which can be 
considered a neurotransmitter. But the adrenal medulla releases epinephrine 
and norepinephrine into circulation, so they should be considered 
hormones. 


What are referred to here as synapses may not fit the strictest definition of 
synapse. Some sources will refer to the connection between a 
postganglionic fiber and a target effector as neuroeffector junctions; 
neurotransmitters, as defined above, would be called neuromodulators. The 
structure of postganglionic connections are not the typical synaptic end bulb 
that is found at the neuromuscular junction, but rather are chains of 
swellings along the length of a postganglionic fiber called a varicosity 
((link]). 

Autonomic Varicosities 


Synaptic vesicles 


> Postganglionic 
varicosities 


Sarcolemma 


The connection between autonomic fibers and target 
effectors is not the same as the typical synapse, such as the 
neuromuscular junction. Instead of a synaptic end bulb, a 
neurotransmitter is released from swellings along the length 
of a fiber that makes an extended network of connections in 
the target effector. 


Note: 

Everyday Connections 

Fight or Flight? What About Fright and Freeze? 

The original usage of the epithet “fight or flight” comes from a scientist 
named Walter Cannon who worked at Harvard in 1915. The concept of 
homeostasis and the functioning of the sympathetic system had been 


introduced in France in the previous century. Cannon expanded the idea, 
and introduced the idea that an animal responds to a threat by preparing to 
stand and fight or run away. The nature of this response was thoroughly 
explained in a book on the physiology of pain, hunger, fear, and rage. 
When students learn about the sympathetic system and the fight-or-flight 
response, they often stop and wonder about other responses. If you were 
faced with a lioness running toward you as pictured at the beginning of this 
chapter, would you run or would you stand your ground? Some people 
would say that they would freeze and not know what to do. So isn’t there 
really more to what the autonomic system does than fight, flight, rest, or 
digest. What about fear and paralysis in the face of a threat? 

The common epithet of “fight or flight” is being enlarged to be “fight, 
flight, or fright” or even “fight, flight, fright, or freeze.” Cannon’s original 
contribution was a catchy phrase to express some of what the nervous 
system does in response to a threat, but it is incomplete. The sympathetic 
system is responsible for the physiological responses to emotional states. 
The name “sympathetic” can be said to mean that (sym- = “together”; - 
pathos = “pain,” “suffering,” or “emotion”). 


Note: 
Oia) 
apr 
= cpenstax couece” 
ony ‘ 
io re 


Watch this video to learn more about the nervous system. As described in 
this video, the nervous system has a way to deal with threats and stress that 
is separate from the conscious control of the somatic nervous system. The 
system comes from a time when threats were about survival, but in the 
modern age, these responses become part of stress and anxiety. This video 
describes how the autonomic system is only part of the response to threats, 
or stressors. What other organ system gets involved, and what part of the 


brain coordinates the two systems for the entire response, including 
epinephrine (adrenaline) and cortisol? 


Chapter Review 


The primary responsibilities of the autonomic nervous system are to 
regulate homeostatic mechanisms in the body, which is also part of what the 
endocrine system does. The key to understanding the autonomic system is 
to explore the response pathways—the output of the nervous system. The 
way we respond to the world around us, to manage the internal environment 
on the basis of the external environment, is divided between two parts of 
the autonomic nervous system. The sympathetic division responds to threats 
and produces a readiness to confront the threat or to run away: the fight-or- 
flight response. The parasympathetic division plays the opposite role. When 
the external environment does not present any immediate danger, a restful 
mode descends on the body, and the digestive system is more active. 


The sympathetic output of the nervous system originates out of the lateral 
horn of the thoracolumbar spinal cord. An axon from one of these central 
neurons projects by way of the ventral spinal nerve root and spinal nerve to 
a sympathetic ganglion, either in the sympathetic chain ganglia or one of 
the collateral locations, where it synapses on a ganglionic neuron. These 
preganglionic fibers release ACh, which excites the ganglionic neuron 
through the nicotinic receptor. The axon from the ganglionic neuron—the 
postganglionic fiber—then projects to a target effector where it will release 
norepinephrine to bind to an adrenergic receptor, causing a change in the 
physiology of that organ in keeping with the broad, divergent sympathetic 
response. The postganglionic connections to sweat glands in the skin and 
blood vessels supplying skeletal muscle are, however, exceptions; those 
fibers release ACh onto muscarinic receptors. The sympathetic system has a 
specialized preganglionic connection to the adrenal medulla that causes 
epinephrine and norepinephrine to be released into the bloodstream rather 
than exciting a neuron that contacts an organ directly. This hormonal 
component means that the sympathetic chemical signal can spread 
throughout the body very quickly and affect many organ systems at once. 


The parasympathetic output is based in the brain stem and sacral spinal 
cord. Neurons from particular nuclei in the brain stem or from the lateral 
horn of the sacral spinal cord (preganglionic neurons) project to terminal 
(intramural) ganglia located close to or within the wall of target effectors. 
These preganglionic fibers also release ACh onto nicotinic receptors to 
excite the ganglionic neurons. The postganglionic fibers then contact the 
target tissues within the organ to release ACh, which binds to muscarinic 
receptors to induce rest-and-digest responses. 


Signaling molecules utilized by the autonomic nervous system are released 
from axons and can be considered as either neurotransmitters (when they 
directly interact with the effector) or as hormones (when they are released 
into the bloodstream). The same molecule, such as norepinephrine, could be 
considered either a neurotransmitter or a hormone on the basis of whether it 
is released from a postganglionic sympathetic axon or from the adrenal 
gland. The synapses in the autonomic system are not always the typical type 
of connection first described in the neuromuscular junction. Instead of 
having synaptic end bulbs at the very end of an axonal fiber, they may have 
swellings—called varicosities—along the length of a fiber so that it makes a 
network of connections within the target tissue. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to learn more about adrenaline and the fight-or-flight 
response. When someone is said to have a rush of adrenaline, the 
image of bungee jumpers or skydivers usually comes to mind. But 
adrenaline, also known as epinephrine, is an important chemical in 
coordinating the body’s fight-or-flight response. In this video, you look 
inside the physiology of the fight-or-flight response, as envisioned for 
a firefighter. His body’s reaction is the result of the sympathetic 
division of the autonomic nervous system causing system-wide 
changes as it prepares for extreme responses. What two changes does 
adrenaline bring about to help the skeletal muscle response? 


Solution: 


The heart rate increases to send more blood to the muscles, and the 
liver releases stored glucose to fuel the muscles. 


Exercise: 


Problem: 


Watch this video to learn more about the nervous system. As described 
in this video, the nervous system has a way to deal with threats and 
stress that is separate from the conscious control of the somatic 
nervous system. The system comes from a time when threats were 
about survival, but in the modern age, these responses become part of 
stress and anxiety. This video describes how the autonomic system is 
only part of the response to threats, or stressors. What other organ 
system gets involved, and what part of the brain coordinates the two 
systems for the entire response, including epinephrine (adrenaline) and 
cortisol? 


Solution: 


The endocrine system is also responsible for responses to stress in our 
lives. The hypothalamus coordinates the autonomic response through 
projections into the spinal cord and through influence over the 
pituitary gland, the effective center of the endocrine system. 


Review Questions 


Exercise: 


Problem: 


Which of these physiological changes would not be considered part of 
the sympathetic fight-or-flight response? 


a. increased heart rate 
b. increased sweating 


c. dilated pupils 
d. increased stomach motility 


Solution: 


D 


Exercise: 


Problem: Which type of fiber could be considered the longest? 


a. preganglionic parasympathetic 
b. preganglionic sympathetic 

c. postganglionic parasympathetic 
d. postganglionic sympathetic 


Solution: 


A 
Exercise: 


Problem: 


Which signaling molecule is most likely responsible for an increase in 
digestive activity? 


a. epinephrine 

b. norepinephrine 
c. acetylcholine 
d. adrenaline 


Solution: 


C 


Exercise: 


Problem: 


Which of these cranial nerves contains preganglionic parasympathetic 
fibers? 


a. optic, CN II 

b. facial, CN VII 

c. trigeminal, CN V 

d. hypoglossal, CN XII 


Solution: 


B 
Exercise: 


Problem: 


Which of the following is not a target of a sympathetic preganglionic 
fiber? 


a. intermural ganglion 
b. collateral ganglion 
c. adrenal gland 

d. chain ganglion 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: 


In the context of a lioness hunting on the savannah, why would the 
sympathetic system not activate the digestive system? 


Solution: 


Whereas energy is needed for running away from the threat, blood 
needs to be sent to the skeletal muscles for oxygen supply. The 
additional fuel, in the form of carbohydrates, probably wouldn’t 
improve the ability to escape the threat as much as the diversion of 
oxygen-rich blood would hinder it. 


Exercise: 


Problem: 


A target effector, such as the heart, receives input from the sympathetic 
and parasympathetic systems. What is the actual difference between 
the sympathetic and parasympathetic divisions at the level of those 
connections (i.e., at the synapse)? 


Solution: 


The postganglionic sympathetic fiber releases norepinephrine, whereas 
the postganglionic parasympathetic fiber releases acetylcholine. 
Specific locations in the heart have adrenergic receptors and 
muscarinic receptors. Which receptors are bound is the signal that 
determines how the heart responds. 


Glossary 
alpha (a)-adrenergic receptor 
one of the receptors to which epinephrine and norepinephrine bind, 


which comes in three subtypes: a, Q>, and a3 


acetylcholine (ACh) 


neurotransmitter that binds at a motor end-plate to trigger 
depolarization 


adrenal medulla 
interior portion of the adrenal (or suprarenal) gland that releases 
epinephrine and norepinephrine into the bloodstream as hormones 


adrenergic 
synapse where norepinephrine is released, which binds to a- or B- 
adrenergic receptors 


beta (B)-adrenergic receptor 
one of the receptors to which epinephrine and norepinephrine bind, 
which comes in two subtypes: 8, and B»5 


celiac ganglion 
one of the collateral ganglia of the sympathetic system that projects to 
the digestive system 


central neuron 
specifically referring to the cell body of a neuron in the autonomic 
system that is located in the central nervous system, specifically the 
lateral horn of the spinal cord or a brain stem nucleus 


cholinergic 
synapse at which acetylcholine is released and binds to the nicotinic or 
muscarinic receptor 


chromaffin cells 
neuroendocrine cells of the adrenal medulla that release epinephrine 
and norepinephrine into the bloodstream as part of sympathetic system 
activity 

ciliary ganglion 
one of the terminal ganglia of the parasympathetic system, located in 


the posterior orbit, axons from which project to the iris 


collateral ganglia 


ganglia outside of the sympathetic chain that are targets of sympathetic 
preganglionic fibers, which are the celiac, inferior mesenteric, and 
superior mesenteric ganglia 


craniosacral system 
alternate name for the parasympathetic division of the autonomic 
nervous system that is based on the anatomical location of central 
neurons in brain-stem nuclei and the lateral horn of the sacral spinal 
cord; also referred to as craniosacral outflow 


dorsal nucleus of the vagus nerve 
location of parasympathetic neurons that project through the vagus 
nerve to terminal ganglia in the thoracic and abdominal cavities 


Eddinger—Westphal nucleus 
location of parasympathetic neurons that project to the ciliary ganglion 


endogenous 
describes substance made in the human body 


epinephrine 
signaling molecule released from the adrenal medulla into the 
bloodstream as part of the sympathetic response 


exogenous 
describes substance made outside of the human body 


fight-or-flight response 
set of responses induced by sympathetic activity that lead to either 
fleeing a threat or standing up to it, which in the modem world is often 
associated with anxious feelings 


G protein—coupled receptor 
membrane protein complex that consists of a receptor protein that 
binds to a signaling molecule—a G protein—that is activated by that 
binding and in tur activates an effector protein (enzyme) that creates a 
second-messenger molecule in the cytoplasm of the target cell 


ganglionic neuron 
specifically refers to the cell body of a neuron in the autonomic system 
that is located in a ganglion 


gray rami Communicantes 
(singular = ramus communicans) unmyelinated structures that provide 
a short connection from a sympathetic chain ganglion to the spinal 
nerve that contains the postganglionic sympathetic fiber 


greater splanchnic nerve 
nerve that contains fibers of the central sympathetic neurons that do 
not synapse in the chain ganglia but project onto the celiac ganglion 


inferior mesenteric ganglion 
one of the collateral ganglia of the sympathetic system that projects to 
the digestive system 


intramural ganglia 
terminal ganglia of the parasympathetic system that are found within 
the walls of the target effector 


lesser splanchnic nerve 
nerve that contains fibers of the central sympathetic neurons that do 
not synapse in the chain ganglia but project onto the inferior 
mesenteric ganglion 


ligand-gated cation channel 
ion channel, such as the nicotinic receptor, that is specific to positively 
charged ions and opens when a molecule such as a neurotransmitter 
binds to it 


mesenteric plexus 
nervous tissue within the wall of the digestive tract that contains 
neurons that are the targets of autonomic preganglionic fibers and that 
project to the smooth muscle and glandular tissues in the digestive 
organ 


muscarinic receptor 


type of acetylcholine receptor protein that is characterized by also 
binding to muscarine and is a metabotropic receptor 


nicotinic receptor 
type of acetylcholine receptor protein that is characterized by also 
binding to nicotine and is an ionotropic receptor 


norepinephrine 
signaling molecule released as a neurotransmitter by most 
postganglionic sympathetic fibers as part of the sympathetic response, 
or as a hormone into the bloodstream from the adrenal medulla 


nucleus ambiguus 
brain-stem nucleus that contains neurons that project through the vagus 
nerve to terminal ganglia in the thoracic cavity; specifically associated 
with the heart 


parasympathetic division 
division of the autonomic nervous system responsible for restful and 
digestive functions 


paravertebral ganglia 
autonomic ganglia superior to the sympathetic chain ganglia 


postganglionic fiber 
axon from a ganglionic neuron in the autonomic nervous system that 
projects to and synapses with the target effector; sometimes referred to 
as a postganglionic neuron 


preganglionic fiber 
axon from a central neuron in the autonomic nervous system that 
projects to and synapses with a ganglionic neuron; sometimes referred 
to as a preganglionic neuron 


prevertebral ganglia 
autonomic ganglia that are anterior to the vertebral column and 
functionally related to the sympathetic chain ganglia 


rest and digest 
set of functions associated with the parasympathetic system that lead 
to restful actions and digestion 


superior cervical ganglion 
one of the paravertebral ganglia of the sympathetic system that 
projects to the head 


superior mesenteric ganglion 
one of the collateral ganglia of the sympathetic system that projects to 
the digestive system 


sympathetic chain ganglia 
series of ganglia adjacent to the vertebral column that receive input 
from central sympathetic neurons 


sympathetic division 
division of the autonomic nervous system associated with the fight-or- 
flight response 


target effector 
organ, tissue, or gland that will respond to the control of an autonomic 
or somatic or endocrine signal 


terminal ganglia 
ganglia of the parasympathetic division of the autonomic system, 
which are located near or within the target effector, the latter also 
known as intramural ganglia 


thoracolumbar system 
alternate name for the sympathetic division of the autonomic nervous 
system that is based on the anatomical location of central neurons in 
the lateral horn of the thoracic and upper lumbar spinal cord 


varicosity 
structure of some autonomic connections that is not a typical synaptic 
end bulb, but a string of swellings along the length of a fiber that 
makes a network of connections with the target effector 


white rami communicantes 
(singular = ramus communicans) myelinated structures that provide a 
short connection from a sympathetic chain ganglion to the spinal nerve 
that contains the preganglionic sympathetic fiber 


Central Control 
By the end of this section, you will be able to: 


¢ Describe the role of higher centers of the brain in autonomic regulation 

e Explain the connection of the hypothalamus to homeostasis 

¢ Describe the regions of the CNS that link the autonomic system with 
emotion 

e Describe the pathways important to descending control of the 
autonomic system 


The pupillary light reflex ((link]) begins when light hits the retina and 
causes a signal to travel along the optic nerve. This is visual sensation, 
because the afferent branch of this reflex is simply sharing the special sense 
pathway. Bright light hitting the retina leads to the parasympathetic 
response, through the oculomotor nerve, followed by the postganglionic 
fiber from the ciliary ganglion, which stimulates the circular fibers of the 
iris to contract and constrict the pupil. When light hits the retina in one eye, 
both pupils contract. When that light is removed, both pupils dilate again 
back to the resting position. When the stimulus is unilateral (presented to 
only one eye), the response is bilateral (both eyes). The same is not true for 
somatic reflexes. If you touch a hot radiator, you only pull that arm back, 
not both. Central control of autonomic reflexes is different than for somatic 
reflexes. The hypothalamus, along with other CNS locations, controls the 
autonomic system. 

Pupillary Reflex Pathways 


Pretectal 


Action potentials 
from right eye reach 
both right and left 
pretectal nuclei. 


Oculomotor SY, VA 
nerves (III) 


Ciliary 
ganglia 
The pretectal 

nuclei stimulate 
both sides of the 
Eddinger—Westphal 
nucleus even 
though the light was 


perceived only in 


() The right and left sides the right eye. 
of the Eddinger—Westphal 


nuclei generate action 


@) Light is shined on 
potentials through the 
right and left oculomotor 


right eye only. 
_<<? 
a 
Cay 
e, nerves, causing both pupils 


Y to constrict. 
A 


The pupil is under competing autonomic control in response 

to light levels hitting the retina. The sympathetic system will 

dilate the pupil when the retina is not receiving enough light, 

and the parasympathetic system will constrict the pupil when 
too much light hits the retina. 


Forebrain Structures 


Autonomic control is based on the visceral reflexes, composed of the 
afferent and efferent branches. These homeostatic mechanisms are based on 
the balance between the two divisions of the autonomic system, which 
results in tone for various organs that is based on the predominant input 
from the sympathetic or parasympathetic systems. Coordinating that 
balance requires integration that begins with forebrain structures like the 
hypothalamus and continues into the brain stem and spinal cord. 


The Hypothalamus 


The hypothalamus is the control center for many homeostatic mechanisms. 
It regulates both autonomic function and endocrine function. The roles it 
plays in the pupillary reflexes demonstrates the importance of this control 
center. The optic nerve projects primarily to the thalamus, which is the 
necessary relay to the occipital cortex for conscious visual perception. 
Another projection of the optic nerve, however, goes to the hypothalamus. 


The hypothalamus then uses this visual system input to drive the pupillary 
reflexes. If the retina is activated by high levels of light, the hypothalamus 
stimulates the parasympathetic response. If the optic nerve message shows 
that low levels of light are falling on the retina, the hypothalamus activates 
the sympathetic response. Output from the hypothalamus follows two main 
tracts, the dorsal longitudinal fasciculus and the medial forebrain bundle 
({link]). Along these two tracts, the hypothalamus can influence the 
Eddinger—Westphal nucleus of the oculomotor complex or the lateral horns 
of the thoracic spinal cord. 

Fiber Tracts of the Central Autonomic System 


Hypothalamus 


Oculomotor cortex 
(includes Eddinger— 
Westphal nucleus) 


Dorsal motor nucleus 
of the vagus 


Nucleus ambiguus 


—— Medial forebrain bundle 
—— Dorsal longitudinal fasciculus 


To spinal cord 


The hypothalamus is the source of most of the central 
control of autonomic function. It receives input from 
cerebral structures and projects to brain stem and 
spinal cord structures to regulate the balance of 
sympathetic and parasympathetic input to the organ 
systems of the body. The main pathways for this are 
the medial forebrain bundle and the dorsal 
longitudinal fasciculus. 


These two tracts connect the hypothalamus with the major parasympathetic 
nuclei in the brain stem and the preganglionic (central) neurons of the 
thoracolumbar spinal cord. The hypothalamus also receives input from 
other areas of the forebrain through the medial forebrain bundle. The 
olfactory cortex, the septal nuclei of the basal forebrain, and the amygdala 
project into the hypothalamus through the medial forebrain bundle. These 
forebrain structures inform the hypothalamus about the state of the nervous 
system and can influence the regulatory processes of homeostasis. A good 
example of this is found in the amygdala, which is found beneath the 
cerebral cortex of the temporal lobe and plays a role in our ability to 
remember and feel emotions. 


The Amygdala 


The amygdala is a group of nuclei in the medial region of the temporal lobe 
that is part of the limbic lobe ({link]). The limbic lobe includes structures 
that are involved in emotional responses, as well as structures that 
contribute to memory function. The limbic lobe has strong connections with 
the hypothalamus and influences the state of its activity on the basis of 
emotional state. For example, when you are anxious or scared, the 
amygdala will send signals to the hypothalamus along the medial forebrain 
bundle that will stimulate the sympathetic fight-or-flight response. The 
hypothalamus will also stimulate the release of stress hormones through its 
control of the endocrine system in response to amygdala input. 

The Limbic Lobe 


Cingulate 


gyrus 
Hypothalamic nuclei 


Corpus 
callosum 


Amygdala 


) : ' SS Et >) bb Ree Thalamus 
Hippocampus . . 


Structures arranged around the edge of the cerebrum 
constitute the limbic lobe, which includes the amygdala, 
hippocampus, and cingulate gyrus, and connects to the 
hypothalamus. 


The Medulla 


The medulla contains nuclei referred to as the cardiovascular center, 
which controls the smooth and cardiac muscle of the cardiovascular system 
through autonomic connections. When the homeostasis of the 
cardiovascular system shifts, such as when blood pressure changes, the 
coordination of the autonomic system can be accomplished within this 
region. Furthermore, when descending inputs from the hypothalamus 
stimulate this area, the sympathetic system can increase activity in the 
cardiovascular system, such as in response to anxiety or stress. The 
preganglionic sympathetic fibers that are responsible for increasing heart 
rate are referred to as the cardiac accelerator nerves, whereas the 
preganglionic sympathetic fibers responsible for constricting blood vessels 
compose the vasomotor nerves. 


Several brain stem nuclei are important for the visceral control of major 
organ systems. One brain stem nucleus involved in cardiovascular function 
is the solitary nucleus. It receives sensory input about blood pressure and 
cardiac function from the glossopharyngeal and vagus nerves, and its output 
will activate sympathetic stimulation of the heart or blood vessels through 
the upper thoracic lateral horn. Another brain stem nucleus important for 
visceral control is the dorsal motor nucleus of the vagus nerve, which is the 
motor nucleus for the parasympathetic functions ascribed to the vagus 
nerve, including decreasing the heart rate, relaxing bronchial tubes in the 
lungs, and activating digestive function through the enteric nervous system. 
The nucleus ambiguus, which is named for its ambiguous histology, also 
contributes to the parasympathetic output of the vagus nerve and targets 
muscles in the pharynx and larynx for swallowing and speech, as well as 
contributing to the parasympathetic tone of the heart along with the dorsal 
motor nucleus of the vagus. 


Note: 

Everyday Connections 

Exercise and the Autonomic System 

In addition to its association with the fight-or-flight response and rest-and- 
digest functions, the autonomic system is responsible for certain everyday 
functions. For example, it comes into play when homeostatic mechanisms 
dynamically change, such as the physiological changes that accompany 
exercise. Getting on the treadmill and putting in a good workout will cause 
the heart rate to increase, breathing to be stronger and deeper, sweat glands 
to activate, and the digestive system to suspend activity. These are the 
same physiological changes associated with the fight-or-flight response, 
but there is nothing chasing you on that treadmill. 

This is not a simple homeostatic mechanism at work because “maintaining 
the internal environment” would mean getting all those changes back to 
their set points. Instead, the sympathetic system has become active during 
exercise so that your body can cope with what is happening. A homeostatic 
mechanism is dealing with the conscious decision to push the body away 
from a resting state. The heart, actually, is moving away from its 
homeostatic set point. Without any input from the autonomic system, the 


heart would beat at approximately 100 bpm, and the parasympathetic 
system slows that down to the resting rate of approximately 70 bpm. But in 
the middle of a good workout, you should see your heart rate at 120-140 
bpm. You could say that the body is stressed because of what you are doing 
to it. Homeostatic mechanisms are trying to keep blood pH in the normal 
range, or to keep body temperature under control, but those are in response 
to the choice to exercise. 


Note: 


Ct [= 


os 
mss" OPENStax COLLEGE 
A “a 
r r. 

io ane 


Watch this video to learn about physical responses to emotion. The 
autonomic system, which is important for regulating the homeostasis of the 
organ systems, is also responsible for our physiological responses to 
emotions such as fear. The video summarizes the extent of the body’s 
reactions and describes several effects of the autonomic system in response 
to fear. On the basis of what you have already studied about autonomic 
function, which effect would you expect to be associated with 
parasympathetic, rather than sympathetic, activity? 


Chapter Review 


The autonomic system integrates sensory information and higher cognitive 
processes to generate output, which balances homeostatic mechanisms. The 
central autonomic structure is the hypothalamus, which coordinates 
sympathetic and parasympathetic efferent pathways to regulate activities of 
the organ systems of the body. The majority of hypothalamic output travels 
through the medial forebrain bundle and the dorsal longitudinal fasciculus 


to influence brain stem and spinal components of the autonomic nervous 
system. The medial forebrain bundle also connects the hypothalamus with 
higher centers of the limbic system where emotion can influence visceral 
responses. The amygdala is a structure within the limbic system that 
influences the hypothalamus in the regulation of the autonomic system, as 
well as the endocrine system. 


These higher centers have descending control of the autonomic system 
through brain stem centers, primarily in the medulla, such as the 
cardiovascular center. This collection of medullary nuclei regulates cardiac 
function, as well as blood pressure. Sensory input from the heart, aorta, and 
carotid sinuses project to these regions of the medulla. The solitary nucleus 
increases sympathetic tone of the cardiovascular system through the cardiac 
accelerator and vasomotor nerves. The nucleus ambiguus and the dorsal 
motor nucleus both contribute fibers to the vagus nerve, which exerts 
parasympathetic control of the heart by decreasing heart rate. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to learn about physical responses to emotion. The 
autonomic system, which is important for regulating the homeostasis 
of the organ systems, is also responsible for our physiological 
responses to emotions such as fear. The video summarizes the extent 
of the body’s reactions and describes several effects of the autonomic 
system in response to fear. On the basis of what you have already 
studied about autonomic function, which effect would you expect to be 
associated with parasympathetic, rather than sympathetic, activity? 


Solution: 


The release of urine in extreme fear. The sympathetic system normally 
constricts sphincters such as that of the urethra. 


Review Questions 


Exercise: 
Problem: 


Which of these locations in the forebrain is the master control center 
for homeostasis through the autonomic and endocrine systems? 


a. hypothalamus 
b. thalamus 

c. amygdala 

d. cerebral cortex 


Solution: 


A 
Exercise: 


Problem: 


Which nerve projects to the hypothalamus to indicate the level of light 
stimuli in the retina? 


a. glossopharyngeal 
b. oculomotor 

c. optic 

d. vagus 


Solution: 


C 


Exercise: 


Problem: 


What region of the limbic lobe is responsible for generating stress 
responses via the hypothalamus? 


a. hippocampus 

b. amygdala 

c. mammillary bodies 
d. prefrontal cortex 


Solution: 


B 
Exercise: 


Problem: 


What is another name for the preganglionic sympathetic fibers that 
project to the heart? 


a. solitary tract 

b. vasomotor nerve 

c. vagus nerve 

d. cardiac accelerator nerve 


Solution: 


D 
Exercise: 


Problem: 


What central fiber tract connects forebrain and brain stem structures 
with the hypothalamus? 


a. cardiac accelerator nerve 


b. medial forebrain bundle 
c. dorsal longitudinal fasciculus 
d. corticospinal tract 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


Horner’s syndrome is a condition that presents with changes in one 
eye, such as pupillary constriction and dropping of eyelids, as well as 
decreased sweating in the face. Why could a tumor in the thoracic 
cavity have an effect on these autonomic functions? 


Solution: 


Pupillary dilation and sweating, two functions lost in Horner’s 
syndrome, are caused by the sympathetic system. A tumor in the 
thoracic cavity may interrupt the output of the thoracic ganglia that 
project to the head and face. 


Exercise: 
Problem: 
The cardiovascular center is responsible for regulating the heart and 
blood vessels through homeostatic mechanisms. What tone does each 
component of the cardiovascular system have? What connections does 


the cardiovascular center invoke to keep these two systems in their 
resting tone? 


Solution: 


The heart—based on the resting heart rate—is under parasympathetic 
tone, and the blood vessels—based on the lack of parasympathetic 
input—are under sympathetic tone. The vagus nerve contributes to the 
lowered resting heart rate, whereas the vasomotor nerves maintain the 
slight constriction of systemic blood vessels. 


Glossary 


cardiac accelerator nerves 
preganglionic sympathetic fibers that cause the heart rate to increase 
when the cardiovascular center in the medulla initiates a signal 


cardiovascular center 
region in the medulla that controls the cardiovascular system through 
cardiac accelerator nerves and vasomotor nerves, which are 
components of the sympathetic division of the autonomic nervous 
system 


dorsal longitudinal fasciculus 
major output pathway of the hypothalamus that descends through the 
gray matter of the brain stem and into the spinal cord 


limbic lobe 
structures arranged around the edges of the cerebrum that are involved 
in memory and emotion 


medial forebrain bundle 
fiber pathway that extends anteriorly into the basal forebrain, passes 
through the hypothalamus, and extends into the brain stem and spinal 
cord 


vasomotor nerves 
preganglionic sympathetic fibers that cause the constriction of blood 
vessels in response to signals from the cardiovascular center 


Organs and Structures of the Respiratory System 
By the end of this section, you will be able to: 


e List the structures that make up the respiratory system 

e Describe how the respiratory system processes oxygen and CO» 

e Compare and contrast the functions of upper respiratory tract with the 
lower respiratory tract 


The major organs of the respiratory system function primarily to provide 
oxygen to body tissues for cellular respiration, remove the waste product 
carbon dioxide, and help to maintain acid-base balance. Portions of the 
respiratory system are also used for non-vital functions, such as sensing 
odors, speech production, and for straining, such as during childbirth or 
coughing ([link]). 

Major Respiratory Structures 


Nasal cavity 
Nostril 


Oral cavit 
J Pharynx 


Larynx 


Trachea 


Left main 
bronchus 


Right main 
bronchus 


Right lun 
: Left lung 


Diaphragm 


The major respiratory structures span the nasal 
cavity to the diaphragm. 


Functionally, the respiratory system can be divided into a conducting zone 
and a respiratory zone. The conducting zone of the respiratory system 
includes the organs and structures not directly involved in gas exchange. 
The gas exchange occurs in the respiratory zone. 


Conducting Zone 


The major functions of the conducting zone are to provide a route for 
incoming and outgoing air, remove debris and pathogens from the incoming 
air, and warm and humidify the incoming air. Several structures within the 
conducting zone perform other functions as well. The epithelium of the 
nasal passages, for example, is essential to sensing odors, and the bronchial 
epithelium that lines the lungs can metabolize some airborne carcinogens. 


The Nose and its Adjacent Structures 


The major entrance and exit for the respiratory system is through the nose. 
When discussing the nose, it is helpful to divide it into two major sections: 
the external nose, and the nasal cavity or internal nose. 


The external nose consists of the surface and skeletal structures that result 
in the outward appearance of the nose and contribute to its numerous 
functions ([link]). The root is the region of the nose located between the 
eyebrows. The bridge is the part of the nose that connects the root to the 
rest of the nose. The dorsum nasi is the length of the nose. The apex is the 
tip of the nose. On either side of the apex, the nostrils are formed by the 
alae (singular = ala). An ala is a cartilaginous structure that forms the 
lateral side of each naris (plural = nares), or nostril opening. The philtrum 
is the concave surface that connects the apex of the nose to the upper lip. 
Nose 


i ee 


Dorsum nasi 


a a as 
———— es _—_ 7" 
i ea Philtrum ew 4 


Frontal bone 


Nasal bone 


Maxillary bone 


Septal cartilage 


Major alar cartilage 


Septal cartilage 


This illustration shows features of the external 
nose (top) and skeletal features of the nose 
(bottom). 


Underneath the thin skin of the nose are its skeletal features (see [link], 
lower illustration). While the root and bridge of the nose consist of bone, 
the protruding portion of the nose is composed of cartilage. As a result, 
when looking at a skull, the nose is missing. The nasal bone is one of a pair 
of bones that lies under the root and bridge of the nose. The nasal bone 
articulates superiorly with the frontal bone and laterally with the maxillary 
bones. Septal cartilage is flexible hyaline cartilage connected to the nasal 
bone, forming the dorsum nasi. The alar cartilage consists of the apex of 
the nose; it surrounds the naris. 


The nares open into the nasal cavity, which is separated into left and right 
sections by the nasal septum ([link]). The nasal septum is formed 
anteriorly by a portion of the septal cartilage (the flexible portion you can 


touch with your fingers) and posteriorly by the perpendicular plate of the 
ethmoid bone (a cranial bone located just posterior to the nasal bones) and 
the thin vomer bones (whose name refers to its plough shape). Each lateral 
wall of the nasal cavity has three bony projections, called the superior, 
middle, and inferior nasal conchae. The inferior conchae are separate bones, 
whereas the superior and middle conchae are portions of the ethmoid bone. 
Conchae serve to increase the surface area of the nasal cavity and to disrupt 
the flow of air as it enters the nose, causing air to bounce along the 
epithelium, where it is cleaned and warmed. The conchae and meatuses 
also conserve water and prevent dehydration of the nasal epithelium by 
trapping water during exhalation. The floor of the nasal cavity is composed 
of the palate. The hard palate at the anterior region of the nasal cavity is 
composed of bone. The soft palate at the posterior portion of the nasal 
cavity consists of muscle tissue. Air exits the nasal cavities via the internal 
nares and moves into the pharynx. 

Upper Airway 

Sphenoidal sinus ee 


Ethmoid bone 


Nasal meatuses Olfactory epithelium 
(superior, middle, 


and inferior) Nasal conchae 


Pharyngeal tonsil Nasal vestibule 


Opening of ase 
auditory tube 
Nasopharynx Hard palate 
Uvula Soft palate 
Tounge 
Palatine tonsil ; 
Lingual tonsil 
se Epiglottis 
Oropharynx 
Hyoid bone 


Laryngopharynx 
Vestibular fold 
Vocal fold 


Thyroid cartilage 


Cricoid cartilage 


Esophagus Thyroid gland 


Trachea 


Several bones that help form the walls of the nasal cavity have air- 
containing spaces called the paranasal sinuses, which serve to warm and 
humidify incoming air. Sinuses are lined with a mucosa. Each paranasal 
sinus is named for its associated bone: frontal sinus, maxillary sinus, 


sphenoidal sinus, and ethmoidal sinus. The sinuses produce mucus and 
lighten the weight of the skull. 


The nares and anterior portion of the nasal cavities are lined with mucous 
membranes, containing sebaceous glands and hair follicles that serve to 
prevent the passage of large debris, such as dirt, through the nasal cavity. 
An olfactory epithelium used to detect odors is found deeper in the nasal 
cavity. 


The conchae, meatuses, and paranasal sinuses are lined by respiratory 
epithelium composed of pseudostratified ciliated columnar epithelium 
({link]). The epithelium contains goblet cells, one of the specialized, 
columnar epithelial cells that produce mucus to trap debris. The cilia of the 
respiratory epithelium help remove the mucus and debris from the nasal 
cavity with a constant beating motion, sweeping materials towards the 
throat to be swallowed. Interestingly, cold air slows the movement of the 
cilia, resulting in accumulation of mucus that may in turn lead to a runny 
nose during cold weather. This moist epithelium functions to warm and 
humidify incoming air. Capillaries located just beneath the nasal epithelium 
warm the air by convection. Serous and mucus-producing cells also secrete 
the lysozyme enzyme and proteins called defensins, which have 
antibacterial properties. Immune cells that patrol the connective tissue deep 
to the respiratory epithelium provide additional protection. 

Pseudostratified Ciliated Columnar Epithelium 


Lumen of 
Goblet cell trachea Cilia 


Pseudostratified 
columnar epithelia 


Seromucous gland 
in submucosa 


Respiratory epithelium is pseudostratified ciliated 
columnar epithelium. Seromucous glands provide 
lubricating mucus. LM x 680. (Micrograph provided by 
the Regents of University of Michigan Medical School © 
2012) 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. 


Pharynx 


The pharynx is a tube formed by skeletal muscle and lined by mucous 
membrane that is continuous with that of the nasal cavities (see [link]). The 
pharynx is divided into three major regions: the nasopharynx, the 
oropharynx, and the laryngopharynx ([link]). 

Divisions of the Pharynx 


Nasal cavity 


Hard palate Soft palate 


Tongue 


Epiglottis 


Larynx (voice box) 


Esophagus 


Trachea 


The pharynx is divided into three regions: the 
nasopharynx, the oropharynx, and the 
laryngopharynx. 


The nasopharynx is flanked by the conchae of the nasal cavity, and it 
serves only as an airway. At the top of the nasopharynx are the pharyngeal 
tonsils. A pharyngeal tonsil, also called an adenoid, is an aggregate of 
lymphoid reticular tissue similar to a lymph node that lies at the superior 
portion of the nasopharynx. The function of the pharyngeal tonsil is not 
well understood, but it contains a rich supply of lymphocytes and is covered 
with ciliated epithelium that traps and destroys invading pathogens that 
enter during inhalation. The pharyngeal tonsils are large in children, but 
interestingly, tend to regress with age and may even disappear. The uvula is 
a small bulbous, teardrop-shaped structure located at the apex of the soft 
palate. Both the uvula and soft palate move like a pendulum during 
swallowing, swinging upward to close off the nasopharynx to prevent 
ingested materials from entering the nasal cavity. In addition, auditory 


(Eustachian) tubes that connect to each middle ear cavity open into the 
nasopharynx. This connection is why colds often lead to ear infections. 


The oropharynx is a passageway for both air and food. The oropharynx is 
bordered superiorly by the nasopharynx and anteriorly by the oral cavity. 
The fauces is the opening at the connection between the oral cavity and the 
oropharynx. As the nasopharynx becomes the oropharynx, the epithelium 
changes from pseudostratified ciliated columnar epithelium to stratified 
squamous epithelium. The oropharynx contains two distinct sets of tonsils, 
the palatine and lingual tonsils. A palatine tonsil is one of a pair of 
structures located laterally in the oropharynx in the area of the fauces. The 
lingual tonsil is located at the base of the tongue. Similar to the pharyngeal 
tonsil, the palatine and lingual tonsils are composed of lymphoid tissue, and 
trap and destroy pathogens entering the body through the oral or nasal 
Cavities. 


The laryngopharynx is inferior to the oropharynx and posterior to the 
larynx. It continues the route for ingested material and air until its inferior 
end, where the digestive and respiratory systems diverge. The stratified 
squamous epithelium of the oropharynx is continuous with the 
laryngopharynx. Anteriorly, the laryngopharynx opens into the larynx, 
whereas posteriorly, it enters the esophagus. 


Larynx 


The larynx is a cartilaginous structure inferior to the laryngopharynx that 
connects the pharynx to the trachea and helps regulate the volume of air that 
enters and leaves the lungs ({link]). The structure of the larynx is formed by 
several pieces of cartilage. Three large cartilage pieces—the thyroid 
cartilage (anterior), epiglottis (superior), and cricoid cartilage (inferior)— 
form the major structure of the larynx. The thyroid cartilage is the largest 
piece of cartilage that makes up the larynx. The thyroid cartilage consists of 
the laryngeal prominence, or “Adam’s apple,” which tends to be more 
prominent in males. The thick cricoid cartilage forms a ring, with a wide 
posterior region and a thinner anterior region. Three smaller, paired 
cartilages—the arytenoids, corniculates, and cuneiforms—attach to the 


epiglottis and the vocal cords and muscle that help move the vocal cords to 
produce speech. 
Larynx 


Epiglottis 
Body of hyoid bone 
Thyrohyoid membrane 
Thyroid cartilage 


Laryngeal prominence 


Cricothyroid ligament 
- 9 Cricoid cartilage 
Cricotracheal ligament 


Tracheal cartilages 


Epiglottis 
Thyrohyoid membrane 
Body of hyoid bone 
Fatty pad 

Thyrohyoid membrane 


Vestibular fold 
Vocal fold 


Thyroid cartilage 
Cricothyroid ligament 


Cuneiform cartilage 
Corniculate cartilage 


Arytenoid cartilage 


Cricoid cartilage 


Cricotracheal ligament 
Tracheal cartilages 


Right lateral view 


The larynx extends from the laryngopharynx and 
the hyoid bone to the trachea. 


The epiglottis, attached to the thyroid cartilage, is a very flexible piece of 
elastic cartilage that covers the opening of the trachea (see [link]). When in 
the “closed” position, the unattached end of the epiglottis rests on the 
glottis. The glottis is composed of the vestibular folds, the true vocal cords, 
and the space between these folds ([link]). A vestibular fold, or false vocal 
cord, is one of a pair of folded sections of mucous membrane. A true vocal 
cord is one of the white, membranous folds attached by muscle to the 
thyroid and arytenoid cartilages of the larynx on their outer edges. The 
inner edges of the true vocal cords are free, allowing oscillation to produce 
sound. The size of the membranous folds of the true vocal cords differs 


between individuals, producing voices with different pitch ranges. Folds in 
males tend to be larger than those in females, which create a deeper voice. 
The act of swallowing causes the pharynx and larynx to lift upward, 
allowing the pharynx to expand and the epiglottis of the larynx to swing 
downward, closing the opening to the trachea. These movements produce a 
larger area for food to pass through, while preventing food and beverages 
from entering the trachea. 

Vocal Cords 


Esophagus Pyriform fossa 
Trachea 
True vocal cord 
Vestibular fold 

Glottis 


Epiglottis 


Tongue 


The true vocal cords and vestibular folds of the 
larynx are viewed inferiorly from the 
laryngopharynx. 


Continuous with the laryngopharynx, the superior portion of the larynx is 
lined with stratified squamous epithelium, transitioning into 
pseudostratified ciliated columnar epithelium that contains goblet cells. 
Similar to the nasal cavity and nasopharynx, this specialized epithelium 
produces mucus to trap debris and pathogens as they enter the trachea. The 
cilia beat the mucus upward towards the laryngopharynx, where it can be 
swallowed down the esophagus. 


Trachea 


The trachea (windpipe) extends from the larynx toward the lungs ((link]a). 
The trachea is formed by 16 to 20 stacked, C-shaped pieces of hyaline 
cartilage that are connected by dense connective tissue. The trachealis 
muscle and elastic connective tissue together form the fibroelastic 
membrane, a flexible membrane that closes the posterior surface of the 
trachea, connecting the C-shaped cartilages. The fibroelastic membrane 
allows the trachea to stretch and expand slightly during inhalation and 
exhalation, whereas the rings of cartilage provide structural support and 
prevent the trachea from collapsing. In addition, the trachealis muscle can 
be contracted to force air through the trachea during exhalation. The trachea 
is lined with pseudostratified ciliated columnar epithelium, which is 
continuous with the larynx. The esophagus borders the trachea posteriorly. 
Trachea 


(a) The tracheal tube is formed by stacked, C-shaped pieces of hyaline 
cartilage. (b) The layer visible in this cross-section of tracheal wall 
tissue between the hyaline cartilage and the lumen of the trachea is the 
mucosa, which is composed of pseudostratified ciliated columnar 
epithelium that contains goblet cells. LM x 1220. (Micrograph 
provided by the Regents of University of Michigan Medical School © 
2012) 


Larynx 


Trachea 
=) Tracheal 
cartilages 


Submucosal Pseudostratified 
seromucous glands columnar epithelia Cilia =©Lumen 


Primary 
bronchi 


Right lung Left lung Hyaline cartilage 


Secondary 
bronchi 


(a) (b) 


Bronchial Tree 


The trachea branches into the right and left primary bronchi at the carina. 
These bronchi are also lined by pseudostratified ciliated columnar 
epithelium containing mucus-producing goblet cells ((link]b). The carina is 
a raised structure that contains specialized nervous tissue that induces 
violent coughing if a foreign body, such as food, is present. Rings of 
cartilage, similar to those of the trachea, support the structure of the bronchi 
and prevent their collapse. The primary bronchi enter the lungs at the hilum, 
a concave region where blood vessels, lymphatic vessels, and nerves also 
enter the lungs. The bronchi continue to branch into bronchial a tree. A 
bronchial tree (or respiratory tree) is the collective term used for these 
multiple-branched bronchi. The main function of the bronchi, like other 
conducting zone structures, is to provide a passageway for air to move into 
and out of each lung. In addition, the mucous membrane traps debris and 
pathogens. 


A bronchiole branches from the tertiary bronchi. Bronchioles, which are 
about 1 mm in diameter, further branch until they become the tiny terminal 
bronchioles, which lead to the structures of gas exchange. There are more 
than 1000 terminal bronchioles in each lung. The muscular walls of the 
bronchioles do not contain cartilage like those of the bronchi. This muscular 
wall can change the size of the tubing to increase or decrease airflow 
through the tube. 


Respiratory Zone 


In contrast to the conducting zone, the respiratory zone includes structures 
that are directly involved in gas exchange. The respiratory zone begins 
where the terminal bronchioles join a respiratory bronchiole, the smallest 
type of bronchiole ({link]), which then leads to an alveolar duct, opening 
into a cluster of alveoli. 

Respiratory Zone 


Terminal bronchiole 


Smooth muscle 


Deoxygenated blood from 
pulmonary artery 


Oxygenated blood to 
pulmonary vein 


Respiratory bronchiole 


Alveolar Alveolus 


sac 


Capillaries 


Alveolar 
pores 


Bronchioles lead to alveolar sacs in the respiratory zone, 
where gas exchange occurs. 


Alveoli 


An alveolar duct is a tube composed of smooth muscle and connective 
tissue, which opens into a cluster of alveoli. An alveolus is one of the many 
small, grape-like sacs that are attached to the alveolar ducts. 


An alveolar sac is a cluster of many individual alveoli that are responsible 
for gas exchange. An alveolus is approximately 200 pm in diameter with 
elastic walls that allow the alveolus to stretch during air intake, which 
greatly increases the surface area available for gas exchange. Alveoli are 
connected to their neighbors by alveolar pores, which help maintain equal 
air pressure throughout the alveoli and lung ([link]). 

Structures of the Respiratory Zone 


(a) The alveolus is responsible for gas exchange. (b) A micrograph 
shows the alveolar structures within lung tissue. LM x 178. 
(Micrograph provided by the Regents of University of Michigan 
Medical School © 2012) 


Alveoli Alveolar duct Blood vessels Lumen of bronchiole 


U> Alveolar pores 
Capillary 
Respiratory membrane * 
Type | alveolar cell 
Macrophage 


Alveolus 
(gas-filled space) 
Type II alveolar cell Alveolar sac 


(a) (b) 


The alveolar wall consists of three major cell types: type I alveolar cells, 
type II alveolar cells, and alveolar macrophages. A type I alveolar cell is a 
squamous epithelial cell of the alveoli, which constitute up to 97 percent of 
the alveolar surface area. These cells are about 25 nm thick and are highly 
permeable to gases. A type II alveolar cell is interspersed among the type I 
cells and secretes pulmonary surfactant, a substance composed of 
phospholipids and proteins that reduces the surface tension of the alveoli. 
Roaming around the alveolar wall is the alveolar macrophage, a 
phagocytic cell of the immune system that removes debris and pathogens 
that have reached the alveoli. 


The simple squamous epithelium formed by type I alveolar cells is attached 
to a thin, elastic basement membrane. This epithelium is extremely thin and 
borders the endothelial membrane of capillaries. Taken together, the alveoli 
and capillary membranes form a respiratory membrane that is 
approximately 0.5 mm thick. The respiratory membrane allows gases to 
cross by simple diffusion, allowing oxygen to be picked up by the blood for 
transport and CO; to be released into the air of the alveoli. 


Note: 

Diseases of the... 

Respiratory System: Asthma 

Asthma is common condition that affects the lungs in both adults and 
children. Approximately 8.2 percent of adults (18.7 million) and 9.4 
percent of children (7 million) in the United States suffer from asthma. In 
addition, asthma is the most frequent cause of hospitalization in children. 
Asthma is a chronic disease characterized by inflammation and edema of 
the airway, and bronchospasms (that is, constriction of the bronchioles), 
which can inhibit air from entering the lungs. In addition, excessive mucus 
secretion can occur, which further contributes to airway occlusion ((link]). 
Cells of the immune system, such as eosinophils and mononuclear cells, 
may also be involved in infiltrating the walls of the bronchi and 
bronchioles. 

Bronchospasms occur periodically and lead to an “asthma attack.” An 
attack may be triggered by environmental factors such as dust, pollen, pet 


hair, or dander, changes in the weather, mold, tobacco smoke, and 
respiratory infections, or by exercise and stress. 
Normal and Bronchial Asthma Tissues 


Mucus 
Goblet cell 
Epithelium 


Basement membrane 


Lamina propria 


O Smooth muscle 
(e) 2 Es - 
y © 6 Gland 


Cartilage 


Mucus 


Goblet cell 
Basement membrane 


Mast cell 
Lamina propria 
Macrophage 
Eosinophil 
Smooth muscle 


Neutrophil 


(a) Normal lung tissue does not have the 
characteristics of lung tissue during (b) an 
asthma attack, which include thickened mucosa, 
increased mucus-producing goblet cells, and 
eosinophil infiltrates. 


Symptoms of an asthma attack involve coughing, shortness of breath, 
wheezing, and tightness of the chest. Symptoms of a severe asthma attack 
that requires immediate medical attention would include difficulty 
breathing that results in blue (cyanotic) lips or face, confusion, drowsiness, 
a rapid pulse, sweating, and severe anxiety. The severity of the condition, 


frequency of attacks, and identified triggers influence the type of 
medication that an individual may require. Longer-term treatments are 
used for those with more severe asthma. Short-term, fast-acting drugs that 
are used to treat an asthma attack are typically administered via an inhaler. 
For young children or individuals who have difficulty using an inhaler, 
asthma medications can be administered via a nebulizer. 

In many cases, the underlying cause of the condition is unknown. 
However, recent research has demonstrated that certain viruses, such as 
human rhinovirus C (HRVC), and the bacteria Mycoplasma pneumoniae 
and Chlamydia pneumoniae that are contracted in infancy or early 
childhood, may contribute to the development of many cases of asthma. 


Note: 
| [a 
= epehstan coulece 
‘ My 
Oe 


Visit this site to learn more about what happens during an asthma attack. 
What are the three changes that occur inside the airways during an asthma 
attack? 


Chapter Review 


The respiratory system is responsible for obtaining oxygen and getting rid 
of carbon dioxide, and aiding in speech production and in sensing odors. 
From a functional perspective, the respiratory system can be divided into 
two major areas: the conducting zone and the respiratory zone. The 
conducting zone consists of all of the structures that provide passageways 
for air to travel into and out of the lungs: the nasal cavity, pharynx, trachea, 
bronchi, and most bronchioles. The nasal passages contain the conchae and 


meatuses that expand the surface area of the cavity, which helps to warm 
and humidify incoming air, while removing debris and pathogens. The 
pharynx is composed of three major sections: the nasopharynx, which is 
continuous with the nasal cavity; the oropharynx, which borders the 
nasopharynx and the oral cavity; and the laryngopharynx, which borders the 
oropharynx, trachea, and esophagus. The respiratory zone includes the 
structures of the lung that are directly involved in gas exchange: the 
terminal bronchioles and alveoli. 


The lining of the conducting zone is composed mostly of pseudostratified 
ciliated columnar epithelium with goblet cells. The mucus traps pathogens 
and debris, whereas beating cilia move the mucus superiorly toward the 
throat, where it is swallowed. As the bronchioles become smaller and 
smaller, and nearer the alveoli, the epithelium thins and is simple squamous 
epithelium in the alveoli. The endothelium of the surrounding capillaries, 
together with the alveolar epithelium, forms the respiratory membrane. This 
is a blood-air barrier through which gas exchange occurs by simple 
diffusion. 


Interactive Link Questions 


Exercise: 


Problem: 


Visit this site to learn more about what happens during an asthma 
attack. What are the three changes that occur inside the airways during 
an asthma attack? 


Solution: 


Inflammation and the production of a thick mucus; constriction of the 
airway muscles, or bronchospasm; and an increased sensitivity to 
allergens. 


Review Questions 


Exercise: 


Problem: 


Which of the following anatomical structures is not part of the 
conducting zone? 


a. pharynx 

b. nasal cavity 
c. alveoli 

d. bronchi 


Solution: 


C 


Exercise: 


Problem: What is the function of the conchae in the nasal cavity? 


a. increase surface area 

b. exchange gases 

c. maintain surface tension 
d. maintain air pressure 


Solution: 


A 
Exercise: 


Problem: 


The fauces connects which of the following structures to the 
oropharynx? 


a. nasopharynx 
b. laryngopharynx 


c. nasal cavity 
d. oral cavity 


Solution: 


D 


Exercise: 


Problem: Which of the following are structural features of the trachea? 


a. C-shaped cartilage 

b. smooth muscle fibers 
c. cilia 

d. all of the above 


Solution: 


A 
Exercise: 


Problem: 


Which of the following structures is not part of the bronchial tree? 


a. alveoli 

b. bronchi 

c. terminal bronchioles 
d. respiratory bronchioles 


Solution: 


C 


Exercise: 


Problem: What is the role of alveolar macrophages? 


a. to secrete pulmonary surfactant 
b. to secrete antimicrobial proteins 
c. to remove pathogens and debris 
d. to facilitate gas exchange 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: Describe the three regions of the pharynx and their functions. 
Solution: 


The pharynx has three major regions. The first region is the 
nasopharynx, which is connected to the posterior nasal cavity and 
functions as an airway. The second region is the oropharynx, which is 
continuous with the nasopharynx and is connected to the oral cavity at 
the fauces. The laryngopharynx is connected to the oropharynx and the 
esophagus and trachea. Both the oropharynx and laryngopharynx are 
passageways for air and food and drink. 


Exercise: 


Problem: 


If a person sustains an injury to the epiglottis, what would be the 
physiological result? 


Solution: 


The epiglottis is a region of the larynx that is important during the 
swallowing of food or drink. As a person swallows, the pharynx moves 
upward and the epiglottis closes over the trachea, preventing food or 
drink from entering the trachea. If a person’s epiglottis were injured, 
this mechanism would be impaired. As a result, the person may have 
problems with food or drink entering the trachea, and possibly, the 
lungs. Over time, this may cause infections such as pneumonia to set 
in. 


Exercise: 


Problem:Compare and contrast the conducting and respiratory zones. 


Solution: 


The conducting zone of the respiratory system includes the organs and 
structures that are not directly involved in gas exchange, but perform 
other duties such as providing a passageway for air, trapping and 
removing debris and pathogens, and warming and humidifying 
incoming air. Such structures include the nasal cavity, pharynx, larynx, 
trachea, and most of the bronchial tree. The respiratory zone includes 
all the organs and structures that are directly involved in gas exchange, 
including the respiratory bronchioles, alveolar ducts, and alveoli. 


References 


Bizzintino J, Lee WM, Laing IA, Vang F, Pappas T, Zhang G, Martin AC, 
Khoo SK, Cox DW, Geelhoed GC, et al. Association between human 
rhinovirus C and severity of acute asthma in children. Eur Respir J 
[Internet]. 2010 [cited 2013 Mar 22]; 37(5):1037—1042. Available from: 


submit=Go&gca=erj%3B37%2F5%2F 1037 &allch= 


Kumar V, Ramzi S, Robbins SL. Robbins Basic Pathology. 7th ed. 
Philadelphia (PA): Elsevier Ltd; 2005. 


Martin RJ, Kraft M, Chu HW, Berns, EA, Cassell GH. A link between 
chronic asthma and chronic infection. J Allergy Clin Immunol [Internet]. 
2001 [cited 2013 Mar 22]; 107(4):595-601. Available from: 


submit=Go&gca=erj%3B37%2F5%2F 1037 &allch= 


Glossary 


ala 
(plural = alae) small, flaring structure of a nostril that forms the lateral 
side of the nares 


alar cartilage 
cartilage that supports the apex of the nose and helps shape the nares; 
it is connected to the septal cartilage and connective tissue of the alae 


alveolar duct 
small tube that leads from the terminal bronchiole to the respiratory 
bronchiole and is the point of attachment for alveoli 


alveolar macrophage 
immune system cell of the alveolus that removes debris and pathogens 


alveolar pore 
opening that allows airflow between neighboring alveoli 


alveolar sac 
cluster of alveoli 


alveolus 
small, grape-like sac that performs gas exchange in the lungs 


apex 
tip of the external nose 


bronchial tree 
collective name for the multiple branches of the bronchi and 
bronchioles of the respiratory system 


bridge 
portion of the external nose that lies in the area of the nasal bones 


bronchiole 
branch of bronchi that are 1 mm or less in diameter and terminate at 
alveolar sacs 


bronchus 
tube connected to the trachea that branches into many subsidiaries and 
provides a passageway for air to enter and leave the lungs 


conducting zone 
region of the respiratory system that includes the organs and structures 
that provide passageways for air and are not directly involved in gas 
exchange 


cricoid cartilage 
portion of the larynx composed of a ring of cartilage with a wide 
posterior region and a thinner anterior region; attached to the 
esophagus 


dorsum nasi 
intermediate portion of the external nose that connects the bridge to the 
apex and is supported by the nasal bone 


epiglottis 
leaf-shaped piece of elastic cartilage that is a portion of the larynx that 
swings to close the trachea during swallowing 


external nose 
region of the nose that is easily visible to others 


fauces 
portion of the posterior oral cavity that connects the oral cavity to the 
oropharynx 


fibroelastic membrane 


specialized membrane that connects the ends of the C-shape cartilage 
in the trachea; contains smooth muscle fibers 


glottis 
opening between the vocal folds through which air passes when 
producing speech 


laryngeal prominence 
region where the two lamina of the thyroid cartilage join, forming a 
protrusion known as “Adam’s apple” 


laryngopharynx 
portion of the pharynx bordered by the oropharynx superiorly and 
esophagus and trachea inferiorly; serves as a route for both air and 
food 


larynx 
cartilaginous structure that produces the voice, prevents food and 
beverages from entering the trachea, and regulates the volume of air 
that enters and leaves the lungs 


lingual tonsil 
lymphoid tissue located at the base of the tongue 


meatus 
one of three recesses (superior, middle, and inferior) in the nasal cavity 
attached to the conchae that increase the surface area of the nasal 
cavity 


naris 
(plural = nares) opening of the nostrils 


nasal bone 
bone of the skull that lies under the root and bridge of the nose and is 
connected to the frontal and maxillary bones 


nasal septum 


wall composed of bone and cartilage that separates the left and right 
nasal cavities 


nasopharynx 
portion of the pharynx flanked by the conchae and oropharynx that 
serves as an airway 


oropharynx 
portion of the pharynx flanked by the nasopharynx, oral cavity, and 
laryngopharynx that is a passageway for both air and food 


palatine tonsil 
one of the paired structures composed of lymphoid tissue located 
anterior to the uvula at the roof of isthmus of the fauces 


paranasal sinus 
one of the cavities within the skull that is connected to the conchae that 
serve to warm and humidify incoming air, produce mucus, and lighten 
the weight of the skull; consists of frontal, maxillary, sphenoidal, and 
ethmoidal sinuses 


pharyngeal tonsil 
structure composed of lymphoid tissue located in the nasopharynx 


pharynx 
region of the conducting zone that forms a tube of skeletal muscle 
lined with respiratory epithelium; located between the nasal conchae 
and the esophagus and trachea 


philtrum 
concave surface of the face that connects the apex of the nose to the 
top lip 

pulmonary surfactant 
substance composed of phospholipids and proteins that reduces the 


surface tension of the alveoli; made by type II alveolar cells 


respiratory bronchiole 


specific type of bronchiole that leads to alveolar sacs 


respiratory epithelium 
ciliated lining of much of the conducting zone that is specialized to 
remove debris and pathogens, and produce mucus 


respiratory membrane 
alveolar and capillary wall together, which form an air-blood barrier 
that facilitates the simple diffusion of gases 


respiratory zone 
includes structures of the respiratory system that are directly involved 
in gas exchange 


root 
region of the external nose between the eyebrows 


thyroid cartilage 
largest piece of cartilage that makes up the larynx and consists of two 
lamina 


trachea 
tube composed of cartilaginous rings and supporting tissue that 
connects the lung bronchi and the larynx; provides a route for air to 
enter and exit the lung 


trachealis muscle 
smooth muscle located in the fibroelastic membrane of the trachea 


true vocal cord 
one of the pair of folded, white membranes that have a free inner edge 
that oscillates as air passes through to produce sound 


type I alveolar cell 
squamous epithelial cells that are the major cell type in the alveolar 
wall; highly permeable to gases 


type II alveolar cell 


cuboidal epithelial cells that are the minor cell type in the alveolar 
wall; secrete pulmonary surfactant 


vestibular fold 
part of the folded region of the glottis composed of mucous membrane; 
supports the epiglottis during swallowing 


The Lungs 
By the end of this section, you will be able to: 


¢ Describe the overall function of the lung 

e Summarize the blood flow pattern associated with the lungs 
¢ Outline the anatomy of the blood supply to the lungs 

e Describe the pleura of the lungs and their function 


A major organ of the respiratory system, each lung houses structures of 
both the conducting and respiratory zones. The main function of the lungs is 
to perform the exchange of oxygen and carbon dioxide with air from the 
atmosphere. To this end, the lungs exchange respiratory gases across a very 
large epithelial surface area—about 70 square meters—that is highly 
permeable to gases. 


Gross Anatomy of the Lungs 


The lungs are pyramid-shaped, paired organs that are connected to the 
trachea by the right and left bronchi; on the inferior surface, the lungs are 
bordered by the diaphragm. The diaphragm is the flat, dome-shaped muscle 
located at the base of the lungs and thoracic cavity. The lungs are enclosed 
by the pleurae, which are attached to the mediastinum. The right lung is 
shorter and wider than the left lung, and the left lung occupies a smaller 
volume than the right. The cardiac notch is an indentation on the surface of 
the left lung, and it allows space for the heart ([link]). The apex of the lung 
is the superior region, whereas the base is the opposite region near the 
diaphragm. The costal surface of the lung borders the ribs. The mediastinal 
surface faces the midline. 

Gross Anatomy of the Lungs 


Trachea 


Superior lobe 


Main (primary) 
bronchus 


Superior lobe Lobar 

(secondary) 
ronchus 

Segmental 


(tertiary) 
bronchus 


Middle lobe Cardiac notch 


Inferior lobe Inferior lobe 


Right lung Left lung 


Each lung is composed of smaller units called lobes. Fissures separate these 
lobes from each other. The right lung consists of three lobes: the superior, 
middle, and inferior lobes. The left lung consists of two lobes: the superior 
and inferior lobes. A bronchopulmonary segment is a division of a lobe, and 
each lobe houses multiple bronchopulmonary segments. Each segment 
receives air from its own tertiary bronchus and is supplied with blood by its 
own artery. Some diseases of the lungs typically affect one or more 
bronchopulmonary segments, and in some cases, the diseased segments can 
be surgically removed with little influence on neighboring segments. A 
pulmonary lobule is a subdivision formed as the bronchi branch into 
bronchioles. Each lobule receives its own large bronchiole that has multiple 
branches. An interlobular septum is a wall, composed of connective tissue, 
which separates lobules from one another. 


Blood Supply and Nervous Innervation of the Lungs 


The blood supply of the lungs plays an important role in gas exchange and 
serves as a transport system for gases throughout the body. In addition, 
innervation by the both the parasympathetic and sympathetic nervous 
systems provides an important level of control through dilation and 
constriction of the airway. 


Blood Supply 


The major function of the lungs is to perform gas exchange, which requires 
blood from the pulmonary circulation. This blood supply contains 
deoxygenated blood and travels to the lungs where erythrocytes, also 
known as red blood cells, pick up oxygen to be transported to tissues 
throughout the body. The pulmonary artery is an artery that arises from 
the pulmonary trunk and carries deoxygenated, arterial blood to the alveoli. 
The pulmonary artery branches multiple times as it follows the bronchi, and 
each branch becomes progressively smaller in diameter. One arteriole and 
an accompanying venule supply and drain one pulmonary lobule. As they 
near the alveoli, the pulmonary arteries become the pulmonary capillary 
network. The pulmonary capillary network consists of tiny vessels with 
very thin walls that lack smooth muscle fibers. The capillaries branch and 
follow the bronchioles and structure of the alveoli. It is at this point that the 
capillary wall meets the alveolar wall, creating the respiratory membrane. 
Once the blood is oxygenated, it drains from the alveoli by way of multiple 
pulmonary veins, which exit the lungs through the hilum. 


Nervous Innervation 


Dilation and constriction of the airway are achieved through nervous 
control by the parasympathetic and sympathetic nervous systems. The 
parasympathetic system causes bronchoconstriction, whereas the 
sympathetic nervous system stimulates bronchodilation. Reflexes such as 
coughing, and the ability of the lungs to regulate oxygen and carbon dioxide 
levels, also result from this autonomic nervous system control. Sensory 
nerve fibers arise from the vagus nerve, and from the second to fifth 
thoracic ganglia. The pulmonary plexus is a region on the lung root 
formed by the entrance of the nerves at the hilum. The nerves then follow 
the bronchi in the lungs and branch to innervate muscle fibers, glands, and 
blood vessels. 


Pleura of the Lungs 


Each lung is enclosed within a cavity that is surrounded by the pleura. The 
pleura (plural = pleurae) is a serous membrane that surrounds the lung. The 
right and left pleurae, which enclose the right and left lungs, respectively, 
are separated by the mediastinum. The pleurae consist of two layers. The 
visceral pleura is the layer that is superficial to the lungs, and extends into 
and lines the lung fissures ({link]). In contrast, the parietal pleura is the 
outer layer that connects to the thoracic wall, the mediastinum, and the 
diaphragm. The visceral and parietal pleurae connect to each other at the 
hilum. The pleural cavity is the space between the visceral and parietal 
layers. 

Parietal and Visceral Pleurae of the Lungs 


Intercostal 
muscle 


Pleural sac | 


Intercostal f , Ke ae 
muscles <4 I ent pleura 


Visceral 
pleura 


Pleural cavity 


Diaphragm 


Chest wall 
(rib cage, sternum, thoracic vertebrae, 
connective tissue, intercostal muscles) 


The pleurae perform two major functions: They produce pleural fluid and 
create cavities that separate the major organs. Pleural fluid is secreted by 
mesothelial cells from both pleural layers and acts to lubricate their 
surfaces. This lubrication reduces friction between the two layers to prevent 
trauma during breathing, and creates surface tension that helps maintain the 
position of the lungs against the thoracic wall. This adhesive characteristic 
of the pleural fluid causes the lungs to enlarge when the thoracic wall 
expands during ventilation, allowing the lungs to fill with air. The pleurae 
also create a division between major organs that prevents interference due 
to the movement of the organs, while preventing the spread of infection. 


Note: 

Everyday Connection 

The Effects of Second-Hand Tobacco Smoke 

The burning of a tobacco cigarette creates multiple chemical compounds 
that are released through mainstream smoke, which is inhaled by the 
smoker, and through sidestream smoke, which is the smoke that is given 
off by the burning cigarette. Second-hand smoke, which is a combination 
of sidestream smoke and the mainstream smoke that is exhaled by the 
smoker, has been demonstrated by numerous scientific studies to cause 
disease. At least 40 chemicals in sidestream smoke have been identified 
that negatively impact human health, leading to the development of cancer 
or other conditions, such as immune system dysfunction, liver toxicity, 
cardiac arrhythmias, pulmonary edema, and neurological dysfunction. 
Furthermore, second-hand smoke has been found to harbor at least 250 
compounds that are known to be toxic, carcinogenic, or both. Some major 
classes of carcinogens in second-hand smoke are polyaromatic 
hydrocarbons (PAHs), N-nitrosamines, aromatic amines, formaldehyde, 
and acetaldehyde. 

Tobacco and second-hand smoke are considered to be carcinogenic. 
Exposure to second-hand smoke can cause lung cancer in individuals who 
are not tobacco users themselves. It is estimated that the risk of developing 
lung cancer is increased by up to 30 percent in nonsmokers who live with 
an individual who smokes in the house, as compared to nonsmokers who 
are not regularly exposed to second-hand smoke. Children are especially 
affected by second-hand smoke. Children who live with an individual who 
smokes inside the home have a larger number of lower respiratory 
infections, which are associated with hospitalizations, and higher risk of 
sudden infant death syndrome (SIDS). Second-hand smoke in the home 
has also been linked to a greater number of ear infections in children, as 
well as worsening symptoms of asthma. 


Chapter Review 


The lungs are the major organs of the respiratory system and are responsible 
for performing gas exchange. The lungs are paired and separated into lobes; 


The left lung consists of two lobes, whereas the right lung consists of three 
lobes. Blood circulation is very important, as blood is required to transport 
oxygen from the lungs to other tissues throughout the body. The function of 
the pulmonary circulation is to aid in gas exchange. The pulmonary artery 
provides deoxygenated blood to the capillaries that form respiratory 
membranes with the alveoli, and the pulmonary veins return newly 
oxygenated blood to the heart for further transport throughout the body. The 
lungs are innervated by the parasympathetic and sympathetic nervous 
systems, which coordinate the bronchodilation and bronchoconstriction of 
the airways. The lungs are enclosed by the pleura, a membrane that is 
composed of visceral and parietal pleural layers. The space between these 
two layers is called the pleural cavity. The mesothelial cells of the pleural 
membrane create pleural fluid, which serves as both a lubricant (to reduce 
friction during breathing) and as an adhesive to adhere the lungs to the 
thoracic wall (to facilitate movement of the lungs during ventilation). 


Review Questions 


Exercise: 


Problem: 
Which of the following structures separates the lung into lobes? 


a. Mediastinum 
b. fissure 

c. root 

d. pleura 


Solution: 


B 


Exercise: 


Problem: 


A section of the lung that receives its own tertiary bronchus is called 
the 


a. bronchopulmonary segment 
b. pulmonary lobule 

c. interpulmonary segment 

d. respiratory segment 


Solution: 


A 
Exercise: 


Problem: 


The circulation picks up oxygen for cellular use and drops 
off carbon dioxide for removal from the body. 


a. pulmonary 
b. interlobular 
c. respiratory 
d. bronchial 


Solution: 


C 
Exercise: 


Problem: 


The pleura that surrounds the lungs consists of two layers, the 


a. visceral and parietal pleurae. 


b. mediastinum and parietal pleurae. 
c. visceral and mediastinum pleurae. 
d. none of the above 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem:Compare and contrast the right and left lungs. 


Solution: 


The right and left lungs differ in size and shape to accommodate other 
organs that encroach on the thoracic region. The right lung consists of 
three lobes and is shorter than the left lung, due to the position of the 
liver underneath it. The left lung consist of two lobes and is longer and 
narrower than the right lung. The left lung has a concave region on the 
mediastinal surface called the cardiac notch that allows space for the 
heart. 


Exercise: 


Problem: Why are the pleurae not damaged during normal breathing? 


Solution: 


There is a cavity, called the pleural cavity, between the parietal and 
visceral layers of the pleura. Mesothelial cells produce and secrete 
pleural fluid into the pleural cavity that acts as a lubricant. Therefore, 
as you breathe, the pleural fluid prevents the two layers of the pleura 
from rubbing against each other and causing damage due to friction. 


Glossary 


bronchoconstriction 
decrease in the size of the bronchiole due to contraction of the 
muscular wall 


bronchodilation 
increase in the size of the bronchiole due to contraction of the 
muscular wall 


cardiac notch 
indentation on the surface of the left lung that allows space for the 
heart 


hilum 
concave structure on the mediastinal surface of the lungs where blood 
vessels, lymphatic vessels, nerves, and a bronchus enter the lung 


lung 
organ of the respiratory system that performs gas exchange 


parietal pleura 
outermost layer of the pleura that connects to the thoracic wall, 
mediastinum, and diaphragm 


pleural cavity 
space between the visceral and parietal pleurae 


pleural fluid 
substance that acts as a lubricant for the visceral and parietal layers of 
the pleura during the movement of breathing 


pulmonary artery 
artery that arises from the pulmonary trunk and carries deoxygenated, 
arterial blood to the alveoli 


pulmonary plexus 


network of autonomic nervous system fibers found near the hilum of 
the lung 


visceral pleura 
innermost layer of the pleura that is superficial to the lungs and 
extends into the lung fissures 


Overview of the Digestive System 
By the end of this section, you will be able to: 


Identify the organs of the alimentary canal from proximal to distal, and 
briefly state their function 

Identify the accessory digestive organs and briefly state their function 
Describe the four fundamental tissue layers of the alimentary canal 
Contrast the contributions of the enteric and autonomic nervous 
systems to digestive system functioning 

Explain how the peritoneum anchors the digestive organs 


The function of the digestive system is to break down the foods you eat, 
release their nutrients, and absorb those nutrients into the body. Although 
the small intestine is the workhorse of the system, where the majority of 
digestion occurs, and where most of the released nutrients are absorbed into 
the blood or lymph, each of the digestive system organs makes a vital 
contribution to this process ([link]). 

Components of the Digestive System 


Salivary glands: 
Mouth Ca Parotid gland 


Tongue 
r Sublingual gland 
Pe Submandibular gland 
Pharynx 


Esophagus 


Liver 


Gallbladder Stomach 


y 7 Spleen 
Small intestine: 


Duodenum 
Jejunum 
lleum 


Pancreas 
Large intestine: 
Transverse colon 


Ascending colon 


Descending colon 
Cecum 

Sigmoid colon 
Appendix 

Rectum 


Anus Anal canal 


All digestive organs play integral roles in 
the life-sustaining process of digestion. 


As is the case with all body systems, the digestive system does not work in 
isolation; it functions cooperatively with the other systems of the body. 
Consider for example, the interrelationship between the digestive and 
cardiovascular systems. Arteries supply the digestive organs with oxygen 
and processed nutrients, and veins drain the digestive tract. These intestinal 
veins, constituting the hepatic portal system, are unique; they do not return 
blood directly to the heart. Rather, this blood is diverted to the liver where 
its nutrients are off-loaded for processing before blood completes its circuit 
back to the heart. At the same time, the digestive system provides nutrients 


to the heart muscle and vascular tissue to support their functioning. The 
interrelationship of the digestive and endocrine systems is also critical. 
Hormones secreted by several endocrine glands, as well as endocrine cells 
of the pancreas, the stomach, and the small intestine, contribute to the 
control of digestion and nutrient metabolism. In turn, the digestive system 
provides the nutrients to fuel endocrine function. [link] gives a quick 
glimpse at how these other systems contribute to the functioning of the 


digestive system. 


Contribution of Other Body Systems to the Digestive System 


Body system 


Cardiovascular 


Endocrine 


Integumentary 


Lymphatic 


Muscular 


Benefits received by the digestive system 


Blood supplies digestive organs with oxygen and 
processed nutrients 


Endocrine hormones help regulate secretion in 
digestive glands and accessory organs 


Skin helps protect digestive organs and 
synthesizes vitamin D for calcium absorption 


Mucosa-associated lymphoid tissue and other 
lymphatic tissue defend against entry of 
pathogens; lacteals absorb lipids; and lymphatic 
vessels transport lipids to bloodstream 


Skeletal muscles support and protect abdominal 
organs 


Contribution of Other Body Systems to the Digestive System 

Body system Benefits received by the digestive system 
Sensory and motor neurons help regulate 

Nervous secretions and muscle contractions in the 


digestive tract 


Respiratory organs provide oxygen and remove 


Respirator aa 
P e carbon dioxide 
Skeletal Bones help protect and support digestive organs 
; Kidneys convert vitamin D into its active form, 
Urinary 


allowing calcium absorption in the small intestine 


Digestive System Organs 


The easiest way to understand the digestive system is to divide its organs 
into two main categories. The first group is the organs that make up the 
alimentary canal. Accessory digestive organs comprise the second group 
and are critical for orchestrating the breakdown of food and the assimilation 
of its nutrients into the body. Accessory digestive organs, despite their 
name, are critical to the function of the digestive system. 


Alimentary Canal Organs 


Also called the gastrointestinal (GI) tract or gut, the alimentary canal 
(aliment- = “to nourish’) is a one-way tube about 7.62 meters (25 feet) in 
length during life and closer to 10.67 meters (35 feet) in length when 
measured after death, once smooth muscle tone is lost. The main function 
of the organs of the alimentary canal is to nourish the body. This tube 
begins at the mouth and terminates at the anus. Between those two points, 
the canal is modified as the pharynx, esophagus, stomach, and small and 


large intestines to fit the functional needs of the body. Both the mouth and 
anus are open to the external environment; thus, food and wastes within the 
alimentary canal are technically considered to be outside the body. Only 
through the process of absorption do the nutrients in food enter into and 


3 cc 


nourish the body’s “inner space.” 


Accessory Structures 


Each accessory digestive organ aids in the breakdown of food ((Link]). 
Within the mouth, the teeth and tongue begin mechanical digestion, 
whereas the salivary glands begin chemical digestion. Once food products 
enter the small intestine, the gallbladder, liver, and pancreas release 
secretions—such as bile and enzymes—essential for digestion to continue. 
Together, these are called accessory organs because they sprout from the 
lining cells of the developing gut (mucosa) and augment its function; 
indeed, you could not live without their vital contributions, and many 
significant diseases result from their malfunction. Even after development 
is complete, they maintain a connection to the gut by way of ducts. 


Histology of the Alimentary Canal 


Throughout its length, the alimentary tract is composed of the same four 
tissue layers; the details of their structural arrangements vary to fit their 
specific functions. Starting from the lumen and moving outwards, these 
layers are the mucosa, submucosa, muscularis, and serosa, which is 
continuous with the mesentery (see [link]). 

Layers of the Alimentary Canal 


Vein 


} 
Submucosal plexus a’, (| | 
(plexus of Meissner) | i Mesentery 


Glands in pelaly 


submucosa <= 


— y Nerve 
Submucosa | y a 


Gland in mucosa 


Duct of gland 
outside tract 


Myenteric plexus 
Lymphatic tissue 
Serosa: 

Areolar connective tissue 
Epithelium 


Lumen 


Mucosa: 
Epithelium 
Lamina propria 
Muscularis mucosae 


Muscularis: 
Circular muscle 
Longitudinal muscle 


The wall of the alimentary canal has four basic tissue 
layers: the mucosa, submucosa, muscularis, and 
serosa. 


The mucosa is referred to as a mucous membrane, because mucus 
production is a characteristic feature of gut epithelium. The membrane 
consists of epithelium, which is in direct contact with ingested food, and the 
lamina propria, a layer of connective tissue analogous to the dermis. In 
addition, the mucosa has a thin, smooth muscle layer, called the muscularis 
mucosa (not to be confused with the muscularis layer, described below). 


Epithelium—lIn the mouth, pharynx, esophagus, and anal canal, the 
epithelium is primarily a non-keratinized, stratified squamous epithelium. 
In the stomach and intestines, it is a simple columnar epithelium. Notice 
that the epithelium is in direct contact with the lumen, the space inside the 
alimentary canal. Interspersed among its epithelial cells are goblet cells, 
which secrete mucus and fluid into the lumen, and enteroendocrine cells, 
which secrete hormones into the interstitial spaces between cells. Epithelial 
cells have a very brief lifespan, averaging from only a couple of days (in the 
mouth) to about a week (in the gut). This process of rapid renewal helps 
preserve the health of the alimentary canal, despite the wear and tear 
resulting from continued contact with foodstuffs. 


Lamina propria—In addition to loose connective tissue, the lamina propria 
contains numerous blood and lymphatic vessels that transport nutrients 
absorbed through the alimentary canal to other parts of the body. The 
lamina propria also serves an immune function by housing clusters of 
lymphocytes, making up the mucosa-associated lymphoid tissue (MALT). 
These lymphocyte clusters are particularly substantial in the distal ileum 
where they are known as Peyer’s patches. When you consider that the 
alimentary canal is exposed to foodborne bacteria and other foreign matter, 
it is not hard to appreciate why the immune system has evolved a means of 
defending against the pathogens encountered within it. 


Muscularis mucosa—This thin layer of smooth muscle is in a constant state 
of tension, pulling the mucosa of the stomach and small intestine into 
undulating folds. These folds dramatically increase the surface area 
available for digestion and absorption. 


As its name implies, the submucosa lies immediately beneath the mucosa. 
A broad layer of dense connective tissue, it connects the overlying mucosa 
to the underlying muscularis. It includes blood and lymphatic vessels 
(which transport absorbed nutrients), and a scattering of submucosal glands 
that release digestive secretions. Additionally, it serves as a conduit for a 
dense branching network of nerves, the submucosal plexus, which functions 
as described below. 


The third layer of the alimentary canal is the muscularis (also called the 
muscularis externa). The muscularis in the small intestine is made up of a 
double layer of smooth muscle: an inner circular layer and an outer 
longitudinal layer. The contractions of these layers promote mechanical 
digestion, expose more of the food to digestive chemicals, and move the 
food along the canal. In the most proximal and distal regions of the 
alimentary canal, including the mouth, pharynx, anterior part of the 
esophagus, and external anal sphincter, the muscularis is made up of 
skeletal muscle, which gives you voluntary control over swallowing and 
defecation. The basic two-layer structure found in the small intestine is 
modified in the organs proximal and distal to it. The stomach is equipped 
for its churning function by the addition of a third layer, the oblique muscle. 
While the colon has two layers like the small intestine, its longitudinal layer 


is segregated into three narrow parallel bands, the tenia coli, which make it 
look like a series of pouches rather than a simple tube. 


The serosa is the portion of the alimentary canal superficial to the 
muscularis. Present only in the region of the alimentary canal within the 
abdominal cavity, it consists of a layer of visceral peritoneum overlying a 
layer of loose connective tissue. Instead of serosa, the mouth, pharynx, and 
esophagus have a dense sheath of collagen fibers called the adventitia. 
These tissues serve to hold the alimentary canal in place near the ventral 
surface of the vertebral column. 


Nerve Supply 


As soon as food enters the mouth, it is detected by receptors that send 
impulses along the sensory neurons of cranial nerves. Without these nerves, 
not only would your food be without taste, but you would also be unable to 
feel either the food or the structures of your mouth, and you would be 
unable to avoid biting yourself as you chew, an action enabled by the motor 
branches of cranial nerves. 


Intrinsic innervation of much of the alimentary canal is provided by the 
enteric nervous system, which runs from the esophagus to the anus, and 
contains approximately 100 million motor, sensory, and interneurons 
(unique to this system compared to all other parts of the peripheral nervous 
system). These enteric neurons are grouped into two plexuses. The 
myenteric plexus (plexus of Auerbach) lies in the muscularis layer of the 
alimentary canal and is responsible for motility, especially the rhythm and 
force of the contractions of the muscularis. The submucosal plexus (plexus 
of Meissner) lies in the submucosal layer and is responsible for regulating 
digestive secretions and reacting to the presence of food (see [link]). 


Extrinsic innervations of the alimentary canal are provided by the 
autonomic nervous system, which includes both sympathetic and 
parasympathetic nerves. In general, sympathetic activation (the fight-or- 
flight response) restricts the activity of enteric neurons, thereby decreasing 
GI secretion and motility. In contrast, parasympathetic activation (the rest- 


and-digest response) increases GI secretion and motility by stimulating 
neurons of the enteric nervous system. 


Blood Supply 


The blood vessels serving the digestive system have two functions. They 
transport the protein and carbohydrate nutrients absorbed by mucosal cells 
after food is digested in the lumen. Lipids are absorbed via lacteals, tiny 
structures of the lymphatic system. The blood vessels’ second function is to 
supply the organs of the alimentary canal with the nutrients and oxygen 
needed to drive their cellular processes. 


Specifically, the more anterior parts of the alimentary canal are supplied 
with blood by arteries branching off the aortic arch and thoracic aorta. 
Below this point, the alimentary canal is supplied with blood by arteries 
branching from the abdominal aorta. The celiac trunk services the liver, 
stomach, and duodenum, whereas the superior and inferior mesenteric 
arteries supply blood to the remaining small and large intestines. 


The veins that collect nutrient-rich blood from the small intestine (where 
most absorption occurs) empty into the hepatic portal system. This venous 
network takes the blood into the liver where the nutrients are either 
processed or stored for later use. Only then does the blood drained from the 
alimentary canal viscera circulate back to the heart. To appreciate just how 
demanding the digestive process is on the cardiovascular system, consider 
that while you are “resting and digesting,” about one-fourth of the blood 
pumped with each heartbeat enters arteries serving the intestines. 


The Peritoneum 


The digestive organs within the abdominal cavity are held in place by the 
peritoneum, a broad serous membranous sac made up of squamous 
epithelial tissue surrounded by connective tissue. It is composed of two 
different regions: the parietal peritoneum, which lines the abdominal wall, 
and the visceral peritoneum, which envelopes the abdominal organs ([link]). 
The peritoneal cavity is the space bounded by the visceral and parietal 


peritoneal surfaces. A few milliliters of watery fluid act as a lubricant to 
minimize friction between the serosal surfaces of the peritoneum. 
The Peritoneum 


Spinal cord 


Vertebra 


Kidney Kidney 


Pancreas Spleen 


Liver Small intestine 


Gallbladder Large intestine 


Large intestine Stomach 


Small intestine 


Visceral peritoneum Peritoneal cavity Parietal peritoneum 
A cross-section of the abdomen shows the 
relationship between abdominal organs and the 


peritoneum (darker lines). 


Note: 

Disorders of the... 

Digestive System: Peritonitis 

Inflammation of the peritoneum is called peritonitis. Chemical peritonitis 
can develop any time the wall of the alimentary canal is breached, allowing 
the contents of the lumen entry into the peritoneal cavity. For example, 
when an ulcer perforates the stomach wall, gastric juices spill into the 
peritoneal cavity. Hemorrhagic peritonitis occurs after a ruptured tubal 
pregnancy or traumatic injury to the liver or spleen fills the peritoneal 
cavity with blood. Even more severe peritonitis is associated with bacterial 
infections seen with appendicitis, colonic diverticulitis, and pelvic 
inflammatory disease (infection of uterine tubes, usually by sexually 


transmitted bacteria). Peritonitis is life threatening and often results in 
emergency surgery to correct the underlying problem and intensive 
antibiotic therapy. When your great grandparents and even your parents 
were young, the mortality from peritonitis was high. Aggressive surgery, 
improvements in anesthesia safety, the advance of critical care expertise, 
and antibiotics have greatly improved the mortality rate from this 
condition. Even so, the mortality rate still ranges from 30 to 40 percent. 


The visceral peritoneum includes multiple large folds that envelope various 
abdominal organs, holding them to the dorsal surface of the body wall. 
Within these folds are blood vessels, lymphatic vessels, and nerves that 
innervate the organs with which they are in contact, supplying their adjacent 
organs. The five major peritoneal folds are described in [link]. Note that 
during fetal development, certain digestive structures, including the first 
portion of the small intestine (called the duodenum), the pancreas, and 
portions of the large intestine (the ascending and descending colon, and the 
rectum) remain completely or partially posterior to the peritoneum. Thus, 
the location of these organs is described as retroperitoneal. 


The Five Major Peritoneal Folds 


Fold Description 
ore Apron-like structure that lies superficial to the small 
intestine and transverse colon; a site of fat deposition 
omentum ; ; 
in people who are overweight 
Falciform Anchors the liver to the anterior abdominal wall and 


ligament inferior border of the diaphragm 


The Five Major Peritoneal Folds 


Fold Description 

heccar Suspends the stomach from the inferior border of the 
liver; provides a pathway for structures connecting to 

omentum 


the liver 


Vertical band of tissue anterior to the lumbar 
Mesentery vertebrae and anchoring all of the small intestine 
except the initial portion (the duodenum) 


Attaches two portions of the large intestine (the 
Mesocolon transverse and sigmoid colon) to the posterior 
abdominal wall 


Note: 


wees Openstax COLLEGE 
oe ee 
- 


By clicking on this link you can watch a short video of what happens to the 
food you eat, as it passes from your mouth to your intestine. Along the 
way, note how the food changes consistency and form. How does this 
change in consistency facilitate your gaining nutrients from food? 


Chapter Review 


The digestive system includes the organs of the alimentary canal and 
accessory structures. The alimentary canal forms a continuous tube that is 


open to the outside environment at both ends. The organs of the alimentary 
canal are the mouth, pharynx, esophagus, stomach, small intestine, and 
large intestine. The accessory digestive structures include the teeth, tongue, 
salivary glands, liver, pancreas, and gallbladder. The wall of the alimentary 
canal is composed of four basic tissue layers: mucosa, submucosa, 
muscularis, and serosa. The enteric nervous system provides intrinsic 
innervation, and the autonomic nervous system provides extrinsic 
innervation. 


Interactive Link Questions 


Exercise: 
Problem: 
By clicking on this link, you can watch a short video of what happens 
to the food you eat as it passes from your mouth to your intestine. 
Along the way, note how the food changes consistency and form. How 


does this change in consistency facilitate your gaining nutrients from 
food? 


Solution: 


Answers may vary. 


Review Questions 


Exercise: 


Problem: 


Which of these organs is not considered an accessory digestive 
structure? 


a. mouth 

b. salivary glands 
c. pancreas 

d. liver 


Solution: 


A 
Exercise: 


Problem: 


Which of the following organs is supported by a layer of adventitia 
rather than serosa? 


a. esophagus 

b. stomach 

c. small intestine 
d. large intestine 


Solution: 
A 
Exercise: 

Problem: Which of the following membranes covers the stomach? 
a. falciform ligament 
b. mesocolon 
c. parietal peritoneum 
d. visceral peritoneum 

Solution: 


D 


Critical Thinking Questions 


Exercise: 
Problem: 
Explain how the enteric nervous system supports the digestive system. 


What might occur that could result in the autonomic nervous system 
having a negative impact on digestion? 


Solution: 


The enteric nervous system helps regulate alimentary canal motility 
and the secretion of digestive juices, thus facilitating digestion. If a 
person becomes overly anxious, sympathetic innervation of the 
alimentary canal is stimulated, which can result in a slowing of 
digestive activity. 


Exercise: 


Problem: 


What layer of the alimentary canal tissue is capable of helping to 
protect the body against disease, and through what mechanism? 


Solution: 


The lamina propria of the mucosa contains lymphoid tissue that makes 
up the MALT and responds to pathogens encountered in the alimentary 
canal. 


Glossary 


accessory digestive organ 
includes teeth, tongue, salivary glands, gallbladder, liver, and pancreas 


alimentary canal 
continuous muscular digestive tube that extends from the mouth to the 
anus 


motility 


movement of food through the GI tract 


mucosa 
innermost lining of the alimentary canal 


muscularis 
muscle (skeletal or smooth) layer of the alimentary canal wall 


myenteric plexus 
(plexus of Auerbach) major nerve supply to alimentary canal wall; 
controls motility 


retroperitoneal 
located posterior to the peritoneum 


serosa 
outermost layer of the alimentary canal wall present in regions within 
the abdominal cavity 


submucosa 
layer of dense connective tissue in the alimentary canal wall that binds 
the overlying mucosa to the underlying muscularis 


submucosal plexus 
(plexus of Meissner) nerve supply that regulates activity of glands and 
smooth muscle 


The Mouth, Pharynx, and Esophagus 
By the end of this section, you will be able to: 


e Describe the structures of the mouth, including its three accessory 
digestive organs 

e Group the 32 adult teeth according to name, location, and function 

e Describe the process of swallowing, including the roles of the tongue, 
upper esophageal sphincter, and epiglottis 

e Trace the pathway food follows from ingestion into the mouth through 
release into the stomach 


In this section, you will examine the anatomy and functions of the three 
main organs of the upper alimentary canal—the mouth, pharynx, and 
esophagus—as well as three associated accessory organs—the tongue, 
salivary glands, and teeth. 


The Mouth 


The cheeks, tongue, and palate frame the mouth, which is also called the 
oral cavity (or buccal cavity). The structures of the mouth are illustrated in 
[link]. 


At the entrance to the mouth are the lips, or labia (singular = labium). Their 
outer covering is skin, which transitions to a mucous membrane in the 
mouth proper. Lips are very vascular with a thin layer of keratin; hence, the 
reason they are "red." They have a huge representation on the cerebral 
cortex, which probably explains the human fascination with kissing! The 
lips cover the orbicularis oris muscle, which regulates what comes in and 
goes out of the mouth. The labial frenulum is a midline fold of mucous 
membrane that attaches the inner surface of each lip to the gum. The cheeks 
make up the oral cavity’s sidewalls. While their outer covering is skin, their 
inner covering is mucous membrane. This membrane is made up of non- 
keratinized, stratified squamous epithelium. Between the skin and mucous 
membranes are connective tissue and buccinator muscles. The next time 
you eat some food, notice how the buccinator muscles in your cheeks and 
the orbicularis oris muscle in your lips contract, helping you keep the food 


from falling out of your mouth. Additionally, notice how these muscles 
work when you are speaking. 


The pocket-like part of the mouth that is framed on the inside by the gums 
and teeth, and on the outside by the cheeks and lips is called the oral 
vestibule. Moving farther into the mouth, the opening between the oral 
cavity and throat (oropharynx) is called the fauces (like the kitchen 
faucet"). The main open area of the mouth, or oral cavity proper, runs from 
the gums and teeth to the fauces. 


When you are chewing, you do not find it difficult to breathe 
simultaneously. The next time you have food in your mouth, notice how the 
arched shape of the roof of your mouth allows you to handle both digestion 
and respiration at the same time. This arch is called the palate. The anterior 
region of the palate serves as a wall (or septum) between the oral and nasal 
cavities as well as a rigid shelf against which the tongue can push food. It is 
created by the maxillary and palatine bones of the skull and, given its bony 
structure, is known as the hard palate. If you run your tongue along the roof 
of your mouth, you’! notice that the hard palate ends in the posterior oral 
cavity, and the tissue becomes fleshier. This part of the palate, known as the 
soft palate, is composed mainly of skeletal muscle. You can therefore 
manipulate, subconsciously, the soft palate—for instance, to yawn, swallow, 
or sing (see [link]). 

Mouth 


Superior lip 
Superior labial frenulum 


wr 
ages AK, Gingivae (gums) 
fg Sy Palatoglossal arch 


Fauces 


Hard palate a 
oe Palatopharyngeal arch 
Soft palate ih 42 
via & [ -\ <0 4 
Palatine tonsil 
Cheek ———————— Vv \ 
Sa ea | 
Tongue (underside) 
IN 
Molars Sa We Lingual frenulum 


~ yee (ix Opening duct of 
Premolars ey ~~) FE py submandibular gland 
y AT) 
: Gingivae (gums) 


Cuspid (canine) we 
Incisors =a ea / 
NS es as, Inferior labial frenulum 
Oral vestibule 
Ne, ee Inferior lip 


Anterior view 


The mouth includes the lips, tongue, palate, gums, and 
teeth. 


A fleshy bead of tissue called the uvula drops down from the center of the 
posterior edge of the soft palate. Although some have suggested that the 
uvula is a vestigial organ, it serves an important purpose. When you 
swallow, the soft palate and uvula move upward, helping to keep foods and 
liquid from entering the nasal cavity. Unfortunately, it can also contribute to 
the sound produced by snoring. Two muscular folds extend downward from 
the soft palate, on either side of the uvula. Toward the front, the 
palatoglossal arch lies next to the base of the tongue; behind it, the 
palatopharyngeal arch forms the superior and lateral margins of the 
fauces. Between these two arches are the palatine tonsils, clusters of 


lymphoid tissue that protect the pharynx. The lingual tonsils are located at 
the base of the tongue. 


The Tongue 


Perhaps you have heard it said that the tongue is the strongest muscle in the 
body. Those who stake this claim cite its strength proportionate to its size. 
Although it is difficult to quantify the relative strength of different muscles, 
it remains indisputable that the tongue is a workhorse, facilitating ingestion, 
mechanical digestion, chemical digestion (lingual lipase), sensation (of 
taste, texture, and temperature of food), swallowing, and vocalization. 


The tongue is attached to the mandible, the styloid processes of the 
temporal bones, and the hyoid bone. The hyoid is unique in that it only 
distantly/indirectly articulates with other bones. The tongue is positioned 
over the floor of the oral cavity. A medial septum extends the entire length 
of the tongue, dividing it into symmetrical halves. 


Beneath its mucous membrane covering, each half of the tongue is 
composed of the same number and type of intrinsic and extrinsic skeletal 
muscles. The intrinsic muscles (those within the tongue) are the 
longitudinalis inferior, longitudinalis superior, transversus linguae, and 
verticalis linguae muscles. These allow you to change the size and shape of 
your tongue, as well as to stick it out, if you wish. Having such a flexible 
tongue facilitates both swallowing and speech. 


As you learned in your study of the muscular system, the extrinsic muscles 
of the tongue are the mylohyoid, hyoglossus, styloglossus, and genioglossus 
muscles. These muscles originate outside the tongue and insert into 
connective tissues within the tongue. The mylohyoid is responsible for 
raising the tongue, the hyoglossus pulls it down and back, the styloglossus 
pulls it up and back, and the genioglossus pulls it forward. Working in 
concert, these muscles perform three important digestive functions in the 
mouth: (1) position food for optimal chewing, (2) gather food into a bolus 
(rounded mass), and (3) position food so it can be swallowed. 


The top and sides of the tongue are studded with papillae, extensions of 
lamina propria of the mucosa, which are covered in stratified squamous 
epithelium ({link]). Fungiform papillae, which are mushroom shaped, cover 
a large area of the tongue; they tend to be larger toward the rear of the 
tongue and smaller on the tip and sides. In contrast, filiform papillae are 
long and thin. Fungiform papillae contain taste buds, and filiform papillae 
have touch receptors that help the tongue move food around in the mouth. 
The filiform papillae create an abrasive surface that performs mechanically, 
much like a cat’s rough tongue that is used for grooming. Lingual glands in 
the lamina propria of the tongue secrete mucus and a watery serous fluid 
that contains the enzyme lingual lipase, which plays a minor role in 
breaking down triglycerides but does not begin working until it is activated 
in the stomach. A fold of mucous membrane on the underside of the tongue, 
the lingual frenulum, tethers the tongue to the floor of the mouth. People 
with the congenital anomaly ankyloglossia, also known by the non-medical 
term “tongue tie,” have a lingual frenulum that is too short or otherwise 
malformed. Severe ankyloglossia can impair speech and must be corrected 
with surgery. 

Tongue 

Epiglottis 


Palatopharyngeal 
arch 


Palatine tonsil 
Lingual tonsil 


Palatoglossal 
arch 


Cg 
<9 
© 


Terminal sulcus 


Foliate papillae 


Circumvallate 
papilla 


Dorsum of tongue 
Fungiform papilla 


Filiform papilla 


This superior view of the tongue 
shows the locations and types of 
lingual papillae. 


The Salivary Glands 


Many small salivary glands are housed within the mucous membranes of 
the mouth and tongue. These minor exocrine glands are constantly secreting 
saliva, either directly into the oral cavity or indirectly through ducts, even 
while you sleep. In fact, an average of 1 to 1.5 liters of saliva is secreted 
each day. Usually just enough saliva is present to moisten the mouth and 
teeth. Secretion increases when you eat, because saliva is essential to 
moisten food and initiate the chemical breakdown of carbohydrates. Small 
amounts of saliva are also secreted by the labial glands in the lips. In 
addition, the buccal glands in the cheeks, palatal glands in the palate, and 
lingual glands in the tongue help ensure that all areas of the mouth are 
supplied with adequate saliva. 


The Major Salivary Glands 


Outside the oral mucosa are three pairs of major salivary glands, which 
secrete the majority of saliva into ducts that open into the mouth: 


e The submandibular glands, which are in the floor of the mouth, 
secrete saliva into the mouth through the submandibular ducts. 

e The sublingual glands, which lie below the tongue, use the lesser 
sublingual ducts to secrete saliva into the oral cavity. 

e The parotid glands lie between the skin and the masseter muscle, near 
the ears. They secrete saliva into the mouth through the parotid duct, 
which is located near the second upper molar tooth ((Link]). 


Saliva 


Saliva is essentially (98 to 99.5 percent) water. The remaining 4.5 percent is 
a complex mixture of ions, glycoproteins, enzymes, growth factors, and 
waste products. Perhaps the most important ingredient in saliva from the 


perspective of digestion is the enzyme salivary amylase, which initiates the 
breakdown of carbohydrates. Food does not spend enough time in the 
mouth to allow all the carbohydrates to break down, but salivary amylase 
continues acting until it is inactivated by stomach acids. Bicarbonate and 
phosphate ions function as chemical buffers, maintaining saliva at a pH 
between 6.35 and 6.85. Salivary mucus helps lubricate food, facilitating 
movement in the mouth, bolus formation, and swallowing. Saliva contains 
immunoglobulin A, which prevents microbes from penetrating the 
epithelium, and lysozyme, which makes saliva antimicrobial. Saliva also 
contains epidermal growth factor, which might have given rise to the adage 
“a mother’s kiss can heal a wound.” 


Each of the major salivary glands secretes a unique formulation of saliva 
according to its cellular makeup. For example, the parotid glands secrete a 
watery solution that contains salivary amylase. The submandibular glands 
have cells similar to those of the parotid glands, as well as mucus-secreting 
cells. Therefore, saliva secreted by the submandibular glands also contains 
amylase but in a liquid thickened with mucus. The sublingual glands 
contain mostly mucous cells, and they secrete the thickest saliva with the 
least amount of salivary amylase. 

Salivary glands 


Parotid salivary gland 


Parotid duct 


Sublingual 
ducts 


Sublingual salivary gland 
Submandibular salivary gland 


Submandibular duct 


The major salivary glands are 
located outside the oral mucosa 


and deliver saliva into the 
mouth through ducts. 


Note: 

Homeostatic Imbalances 

The Parotid Glands: Mumps 

Infections of the nasal passages and pharynx can attack any salivary gland. 
The parotid glands are the usual site of infection with the virus that causes 
mumps (paramyxovirus). Mumps manifests by enlargement and 
inflammation of the parotid glands, causing a characteristic swelling 
between the ears and the jaw. Symptoms include fever and throat pain, 
which can be severe when swallowing acidic substances such as orange 
juice. 

In about one-third of men who are past puberty, mumps also causes 
testicular inflammation, typically affecting only one testis and rarely 
resulting in sterility. With the increasing use and effectiveness of mumps 
vaccines, the incidence of mumps has decreased dramatically. According to 
the U.S. Centers for Disease Control and Prevention (CDC), the number of 
mumps cases dropped from more than 150,000 in 1968 to fewer than 1700 
in 1993 to only 11 reported cases in 2011. 


Regulation of Salivation 


The autonomic nervous system regulates salivation (the secretion of 
Saliva). In the absence of food, parasympathetic stimulation keeps saliva 
flowing at just the right level for comfort as you speak, swallow, sleep, and 
generally go about life. Over-salivation can occur, for example, if you are 
stimulated by the smell of food, but that food is not available for you to eat. 
Drooling is an extreme instance of the overproduction of saliva. During 
times of stress, such as before speaking in public, sympathetic stimulation 
takes over, reducing salivation and producing the symptom of dry mouth 
often associated with anxiety. When you are dehydrated, salivation is 


reduced, causing the mouth to feel dry and prompting you to take action to 
quench your thirst. 


Salivation can be stimulated by the sight, smell, and taste of food. It can 
even be stimulated by thinking about food. You might notice whether 
reading about food and salivation right now has had any effect on your 
production of saliva. 


How does the salivation process work while you are eating? Food contains 
chemicals that stimulate taste receptors on the tongue, which send impulses 
to the superior and inferior salivatory nuclei in the brain stem. These two 
nuclei then send back parasympathetic impulses through fibers in the 
glossopharyngeal and facial nerves, which stimulate salivation. Even after 
you swallow food, salivation is increased to cleanse the mouth and to water 
down and neutralize any irritating chemical remnants, such as that hot sauce 
in your burrito. Most saliva is swallowed along with food and is reabsorbed, 
so that fluid is not lost. 


The Teeth 


The teeth, or dentes (singular = dens), are organs similar to bones that you 
use to tear, grind, and otherwise mechanically break down food. 


Types of Teeth 


During the course of your lifetime, you have two sets of teeth (one set of 
teeth is a dentition). Your 20 deciduous teeth, or baby teeth, first begin to 
appear at about 6 months of age. Between approximately age 6 and 12, 
these teeth are replaced by 32 permanent teeth. Moving from the center of 
the mouth toward the side, these are as follows ([link]): 


e The eight incisors, four top and four bottom, are the sharp front teeth 
you use for biting into food. 

e The four cuspids (or canines) flank the incisors and have a pointed 
edge (cusp) to tear up food. These fang-like teeth are superb for 
piercing tough or fleshy foods. 


e Posterior to the cuspids are the eight premolars (or bicuspids), which 
have an overall flatter shape with two rounded cusps useful for 
mashing foods. 

e The most posterior and largest are the 12 molars, which have several 
pointed cusps used to crush food so it is ready for swallowing. The 
third members of each set of three molars, top and bottom, are 
commonly referred to as the wisdom teeth, because their eruption is 
commonly delayed until early adulthood. It is not uncommon for 
wisdom teeth to fail to erupt; that is, they remain impacted. In these 
cases, the teeth are typically removed by orthodontic surgery. 


Permanent and Deciduous Teeth 


Central incisor (7-8 yr) 
Lateral incisor (8-9 yr) 
Cuspid or canine (11-12 yr) 
First premolar or 

bicuspid (9-10 yr) 

Second premolar or 
bicuspid (10—12 yr) 


First molar (6-7 yr) 


Second molar 
(12-13 yr) 


Third molar or 
Central incisor wisdom tooth 
(8-12 mo) 


Lateral incisor (12-24 mo) 
Cuspid or canine 
(16-24 mo) 


First molar (12-16 mo) 


Second molar (24-32 mo) 


Second molar (24-32 mo) 


First molar (12-16 mo) 


Cuspid or canine 
(16-24 mo) 
Lateral incisor (12-15 mo) 


Central incisor 
(6-8 mo) Third molar or 


wisdom tooth 


Second molar 
(11-13 yr) 


First molar (6-7 yr) 
Second premolar or 
bicuspid (11-12 yr) 

First premolar or 

bicuspid (9-10 yr) 

Cuspid or canine (9-10 yr) 
Lateral incisor (7-8 yr) 


Central incisor (7-8 yr) 


This figure of two human dentitions 
shows the arrangement of teeth in the 
maxilla and mandible, and the 
relationship between the deciduous 
and permanent teeth. 


Anatomy of a Tooth 


The teeth are secured in the alveolar processes (sockets) of the maxilla and 
the mandible. Gingivae (commonly called the gums) are soft tissues that 
line the alveolar processes and surround the necks of the teeth. Teeth are 
also held in their sockets by a connective tissue called the periodontal 
ligament. 


The two main parts of a tooth are the crown, which is the portion projecting 
above the gum line, and the root, which is embedded within the maxilla and 
mandible. Both parts contain an inner pulp cavity, containing loose 
connective tissue through which run nerves and blood vessels. The region 
of the pulp cavity that runs through the root of the tooth is called the root 
canal. Surrounding the pulp cavity is dentin, a bone-like tissue. In the root 
of each tooth, the dentin is covered by an even harder bone-like layer called 
cementum. In the crown of each tooth, the dentin is covered by an outer 
layer of enamel, the hardest substance in the body ([link]). 


Although enamel protects the underlying dentin and pulp cavity, it is still 
nonetheless susceptible to mechanical and chemical erosion, or what is 
known as tooth decay. The most common form, dental caries (cavities) 
develops when colonies of bacteria feeding on sugars in the mouth release 
acids that cause soft tissue inflammation and degradation of the calcium 
crystals of the enamel. The digestive functions of the mouth are 
summarized in [link]. 

The Structure of the Tooth 


Enamel 


crown Dentin 


—— Gingiva 


Neck _ \ (gum) 

\ 

\ ‘ 
=m Pulp cavity 
~; & \ (contains 
| |g blood vessels 

TD tt V&, 
Lf ') 7S and nerves) 
16 i a | (0 
Root (| ad l ime. Periodontal 
NL) i » ee ligament 
\\ “ Lt] f : 
HO {| 771 Root canal 
|G, / > Ae 
| V4 I . \ 
WM! SS 
Lp ey 26 7 — Bone 


This longitudinal section through a 


molar in its alveolar socket shows the 
relationships between enamel, dentin, 


Digestive Functions of the Mouth 


Structure 


Lips and 
cheeks 


Salivary 
glands 


Tongue’s 
extrinsic 
muscles 


and pulp. 


Action 


Confine 
food 
between 
teeth 


Secrete 
saliva 


Move 
tongue 


sideways, 


and in 
and out 


Outcome 


Food is chewed evenly during 
mastication 


Moisten and lubricate the lining 
of the mouth and pharynx 
Moisten, soften, and dissolve food 
Clean the mouth and teeth 
Salivary amylase breaks down 
starch 


Manipulate food for chewing 
Shape food into a bolus 
Manipulate food for swallowing 


Digestive Functions of the Mouth 


Structure Action Outcome 
Tongue’s Change 
intrinsic tongue Manipulate food for swallowing 
muscles shape 
Sense Nerve impulses from taste buds 
T food in are conducted to salivary nuclei in 
aste 
mouth the brain stem and then to salivary 
buds é 
and sense glands, stimulating saliva 
taste secretion 
; Secrete Activated in the stomach 
Lingual ; ; eager 
fande lingual Break down triglycerides into 
s lipase fatty acids and diglycerides 
Sales Break down solid food into 
Teeth and crush - 
smaller particles for deglutition 
food 
The Pharynx 


The pharynx (throat) is involved in both digestion and respiration. It 
receives food and air from the mouth, and air from the nasal cavities. When 
food enters the pharynx, involuntary muscle contractions close off the air 
passageways. 


A short tube of skeletal muscle lined with a mucous membrane, the pharynx 
runs from the posterior oral and nasal cavities to the opening of the 
esophagus and larynx. It has three subdivisions. The most superior, the 


nasopharynx, is involved only in breathing and speech. The other two 
subdivisions, the oropharynx and the laryngopharynx, are used for both 
breathing and digestion. The oropharynx begins inferior to the nasopharynx 
and is continuous below with the laryngopharynx ([link]). The inferior 
border of the laryngopharynx connects to the esophagus, whereas the 
anterior portion connects to the larynx, allowing air to flow into the 
bronchial tree. 

Pharynx 


Soft palate 


Nasopharynx 
Hard palate 


Uvula 


Oropharynx 

Epiglottis 
Glottis \ Laryngopharynx 
Larynx 
Trachea 
—~ 

Esophagus 
Es Nasal cavity 
| Oral cavity S) —/ 
= Pharynx 


aa Larynx a ae — 


The pharynx runs from the nostrils to the 
esophagus and the larynx. 


Histologically, the wall of the oropharynx is similar to that of the oral 
cavity. The mucosa includes a stratified squamous epithelium that is 
endowed with mucus-producing glands. During swallowing, the elevator 
skeletal muscles of the pharynx contract, raising and expanding the pharynx 


to receive the bolus of food. Once received, these muscles relax and the 
constrictor muscles of the pharynx contract, forcing the bolus into the 
esophagus and initiating peristalsis. 


Usually during swallowing, the soft palate and uvula rise reflexively to 
close off the entrance to the nasopharynx. At the same time, the larynx is 
pulled superiorly and the cartilaginous epiglottis, its most superior structure, 
folds inferiorly, covering the glottis (the opening to the larynx); this process 
effectively blocks access to the trachea and bronchi. When the food “goes 
down the wrong way,” it goes into the trachea. When food enters the 
trachea, the reaction is to cough, which usually forces the food up and out 
of the trachea, and back into the pharynx. 


The Esophagus 


The esophagus is a muscular tube that connects the pharynx to the stomach. 
It is approximately 25.4 cm (10 in) in length, located posterior to the 
trachea, and remains in a collapsed form when not engaged in swallowing. 
As you can see in [link], the esophagus runs a mainly straight route through 
the mediastinum of the thorax. To enter the abdomen, the esophagus 
penetrates the diaphragm through an opening called the esophageal hiatus. 


Passage of Food through the Esophagus 


The upper esophageal sphincter, which is continuous with the inferior 
pharyngeal constrictor, controls the movement of food from the pharynx 
into the esophagus. The upper two-thirds of the esophagus consists of both 
smooth and skeletal muscle fibers, with the latter fading out in the bottom 
third of the esophagus. Rhythmic waves of peristalsis, which begin in the 
upper esophagus, propel the bolus of food toward the stomach. Meanwhile, 
secretions from the esophageal mucosa lubricate the esophagus and food. 
Food passes from the esophagus into the stomach at the lower esophageal 
sphincter (also called the gastroesophageal or cardiac sphincter). Recall 
that sphincters are muscles that surround tubes and serve as valves, closing 
the tube when the sphincters contract and opening it when they relax. The 


lower esophageal sphincter relaxes to let food pass into the stomach, and 
then contracts to prevent stomach acids from backing up into the 
esophagus. Surrounding this sphincter is the muscular diaphragm, which 
helps close off the sphincter when no food is being swallowed. When the 
lower esophageal sphincter does not completely close, the stomach’s 
contents can reflux (that is, back up into the esophagus), causing heartburn 
or gastroesophageal reflux disease (GERD). 

Esophagus 


Upper esophageal 
sphincter 


Esophagus Lower esophageal 


sphincter 
{8) 


Stomach SS 
TY 


The upper esophageal sphincter 
controls the movement of food 
from the pharynx to the 
esophagus. The lower esophageal 
sphincter controls the movement 
of food from the esophagus to the 
stomach. 


Histology of the Esophagus 


The mucosa of the esophagus is made up of an epithelial lining that 
contains non-keratinized, stratified squamous epithelium, with a layer of 
basal and parabasal cells. This epithelium protects against erosion from 
food particles. The mucosa’s lamina propria contains mucus-secreting 
glands. The muscularis layer changes according to location: In the upper 
third of the esophagus, the muscularis is skeletal muscle. In the middle 
third, it is both skeletal and smooth muscle. In the lower third, it is smooth 
muscle. As mentioned previously, the most superficial layer of the 
esophagus is called the adventitia, not the serosa. In contrast to the stomach 
and intestines, the loose connective tissue of the adventitia is not covered by 
a fold of visceral peritoneum. The digestive functions of the esophagus are 
identified in [link]. 


Digestive Functions of the Esophagus 


Action Outcome 

Upper 

esophageal Allows the bolus to move from the 

sphincter laryngopharynx to the esophagus 

relaxation 

Peristalsis Propels the bolus through the esophagus 
powel Allows the bolus to move from the esophagus 
esophageal 


into the stomach and prevents chime from 


hincter ' 
ee entering the esophagus 


relaxation 


Digestive Functions of the Esophagus 


Action Outcome 
Mucus Lubricates the esophagus, allowing easy passage 
secretion of the bolus 

Deglutition 


Deglutition is another word for swallowing—the movement of food from 
the mouth to the stomach. The entire process takes about 4 to 8 seconds for 
solid or semisolid food, and about 1 second for very soft food and liquids. 
Although this sounds quick and effortless, deglutition is, in fact, a complex 
process that involves both the skeletal muscle of the tongue and the muscles 
of the pharynx and esophagus. It is aided by the presence of mucus and 
saliva. There are three stages in deglutition: the voluntary phase, the 
pharyngeal phase, and the esophageal phase ([link]). The autonomic 
nervous system controls the latter two phases. 

Deglutition 


je = “aS 
Y Kg 
HSS 
bn. m 
_— < ¢ 
} Ino 
) 


Superior 
pharyngeal 
constrictor 
muscle 


Medial 
pharyngeal 
constrictor 
muscle 


Medial and 

inferior 

\ i pharyngeal 
\@\\ constrictor 

' \ muscles . 

Inferior 


pharyngeal 

| and esophageal 
\\\—— constrictor 
muscles 


<<“ 
“a 

\ 

A 

( 

\ 

YW 

Yj 


Deglutition includes the voluntary phase and two 
involuntary phases: the pharyngeal phase and the 
esophageal phase. 


The Voluntary Phase 


The voluntary phase of deglutition (also known as the oral or buccal 
phase) is so called because you can control when you swallow food. In this 
phase, chewing has been completed and swallowing is set in motion. The 
tongue moves upward and backward against the palate, pushing the bolus to 
the back of the oral cavity and into the oropharynx. Other muscles keep the 
mouth closed and prevent food from falling out. At this point, the two 
involuntary phases of swallowing begin. 


The Pharyngeal Phase 


In the pharyngeal phase, stimulation of receptors in the oropharynx sends 
impulses to the deglutition center (a collection of neurons that controls 
swallowing) in the medulla oblongata. Impulses are then sent back to the 
uvula and soft palate, causing them to move upward and close off the 
nasopharynx. The laryngeal muscles also constrict to prevent aspiration of 
food into the trachea. At this point, deglutition apnea takes place, which 
means that breathing ceases for a very brief time. Contractions of the 
pharyngeal constrictor muscles move the bolus through the oropharynx and 
laryngopharynx. Relaxation of the upper esophageal sphincter then allows 
food to enter the esophagus. 


The Esophageal Phase 


The entry of food into the esophagus marks the beginning of the esophageal 
phase of deglutition and the initiation of peristalsis. As in the previous 
phase, the complex neuromuscular actions are controlled by the medulla 
oblongata. Peristalsis propels the bolus through the esophagus and toward 
the stomach. The circular muscle layer of the muscularis contracts, pinching 


the esophageal wall and forcing the bolus forward. At the same time, the 
longitudinal muscle layer of the muscularis also contracts, shortening this 
area and pushing out its walls to receive the bolus. In this way, a series of 
contractions keeps moving food toward the stomach. When the bolus nears 
the stomach, distention of the esophagus initiates a short reflex relaxation of 
the lower esophageal sphincter that allows the bolus to pass into the 
stomach. During the esophageal phase, esophageal glands secrete mucus 
that lubricates the bolus and minimizes friction. 


Note: 


I 


OR ero 
HS Erk ep 
wees OPenstax COLLEGE” 


one 


Watch this_animation to see how swallowing is a complex process that 
involves the nervous system to coordinate the actions of upper respiratory 
and digestive activities. During which stage of swallowing is there a risk of 
food entering respiratory pathways and how is this risk blocked? 


Chapter Review 


In the mouth, the tongue and the teeth begin mechanical digestion, and 
saliva begins chemical digestion. The pharynx, which plays roles in 
breathing and vocalization as well as digestion, runs from the nasal and oral 
cavities superiorly to the esophagus inferiorly (for digestion) and to the 
larynx anteriorly (for respiration). During deglutition (swallowing), the soft 
palate rises to close off the nasopharynx, the larynx elevates, and the 
epiglottis folds over the glottis. The esophagus includes an upper 
esophageal sphincter made of skeletal muscle, which regulates the 
movement of food from the pharynx to the esophagus. It also has a lower 
esophageal sphincter, made of smooth muscle, which controls the passage 


of food from the esophagus to the stomach. Cells in the esophageal wall 
secrete mucus that eases the passage of the food bolus. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this animation to see how swallowing is a complex process that 
involves the nervous system to coordinate the actions of upper 
respiratory and digestive activities. During which stage of swallowing 
is there a risk of food entering respiratory pathways and how is this 
risk blocked? 


Solution: 


Answers may vary. 


Review Questions 


Exercise: 


Problem: 


Which of these ingredients in saliva is responsible for activating 
salivary amylase? 


a. mucus 

b. phosphate ions 
c. chloride ions 
d. urea 


Solution: 


C 


Exercise: 


Problem: Which of these statements about the pharynx is true? 


a. It extends from the nasal and oral cavities superiorly to the 
esophagus anteriorly. 

b. The oropharynx is continuous superiorly with the nasopharynx. 

c. The nasopharynx is involved in digestion. 

d. The laryngopharynx is composed partially of cartilage. 


Solution: 


B 
Exercise: 


Problem: 


Which structure is located where the esophagus penetrates the 
diaphragm? 


a. esophageal hiatus 

b. cardiac orifice 

c. upper esophageal sphincter 
d. lower esophageal sphincter 


Solution: 


A 
Exercise: 


Problem: 


Which phase of deglutition involves contraction of the longitudinal 
muscle layer of the muscularis? 


a. voluntary phase 


b. buccal phase 
c. pharyngeal phase 
d. esophageal phase 


Solution: 


D 


Critical Thinking Questions 


Exercise: 


Problem: 


The composition of saliva varies from gland to gland. Discuss how 
saliva produced by the parotid gland differs in action from saliva 
produced by the sublingual gland. 


Solution: 


Parotid gland saliva is watery with little mucus but a lot of amylase, 
which allows it to mix freely with food during mastication and begin 
the digestion of carbohydrates. In contrast, sublingual gland saliva has 
a lot of mucus with the least amount of amylase of all the salivary 
glands. The high mucus content serves to lubricate the food for 
swallowing. 


Exercise: 
Problem: 
During a hockey game, the puck hits a player in the mouth, knocking 


out all eight of his most anterior teeth. Which teeth did the player lose 
and how does this loss affect food ingestion? 


Solution: 


The incisors. Since these teeth are used for tearing off pieces of food 
during ingestion, the player will need to ingest foods that have already 
been cut into bite-sized pieces until the broken teeth are replaced. 


Exercise: 


Problem: What prevents swallowed food from entering the airways? 
Solution: 


Usually when food is swallowed, involuntary muscle contractions 
cause the soft palate to rise and close off the nasopharynx. The larynx 
also is pulled up, and the epiglottis folds over the glottis. These actions 
block off the air passages. 


Exercise: 


Problem: 
Explain the mechanism responsible for gastroesophageal reflux. 
Solution: 


If the lower esophageal sphincter does not close completely, the 
stomach’s acidic contents can back up into the esophagus, a 
phenomenon known as GERD. 


Exercise: 
Problem: 


Describe the three processes involved in the esophageal phase of 
deglutition. 


Solution: 
Peristalsis moves the bolus down the esophagus and toward the 


stomach. Esophageal glands secrete mucus that lubricates the bolus 
and reduces friction. When the bolus nears the stomach, the lower 


esophageal sphincter relaxes, allowing the bolus to pass into the 
stomach. 


References 


van Loon FPL, Holmes SJ, Sirotkin B, Williams W, Cochi S, Hadler S, 
Lindegren ML. Morbidity and Mortality Weekly Report: Mumps 
surveillance -- United States, 1988—1993 [Internet]. Atlanta, GA: Center for 
Disease Control; [cited 2013 Apr 3]. Available from: 
http://www.cdc.gov/mmwr/preview/mmwrhtml/00038546.htm. 


Glossary 


bolus 
mass of chewed food 


cementum 
bone-like tissue covering the root of a tooth 


crown 
portion of tooth visible superior to the gum line 


cuspid 
(also, canine) pointed tooth used for tearing and shredding food 


deciduous tooth 
one of 20 “baby teeth” 


deglutition 
three-stage process of swallowing 


dens 
tooth 


dentin 
bone-like tissue immediately deep to the enamel of the crown or 
cementum of the root of a tooth 


dentition 
set of teeth 


enamel 
covering of the dentin of the crown of a tooth 


esophagus 
muscular tube that runs from the pharynx to the stomach 


fauces 
opening between the oral cavity and the oropharynx 


gingiva 
gum 


incisor 
midline, chisel-shaped tooth used for cutting into food 


labium 
lip 


labial frenulum 
midline mucous membrane fold that attaches the inner surface of the 
lips to the gums 


laryngopharynx 
part of the pharynx that functions in respiration and digestion 


lingual frenulum 
mucous membrane fold that attaches the bottom of the tongue to the 
floor of the mouth 


lingual lipase 
digestive enzyme from glands in the tongue that acts on triglycerides 


lower esophageal sphincter 
smooth muscle sphincter that regulates food movement from the 
esophagus to the stomach 


molar 
tooth used for crushing and grinding food 


oral cavity 
(also, buccal cavity) mouth 


oral vestibule 
part of the mouth bounded externally by the cheeks and lips, and 
internally by the gums and teeth 


oropharynx 
part of the pharynx continuous with the oral cavity that functions in 
respiration and digestion 


palatoglossal arch 
muscular fold that extends from the lateral side of the soft palate to the 
base of the tongue 


palatopharyngeal arch 
muscular fold that extends from the lateral side of the soft palate to the 
side of the pharynx 


parotid gland 
one of a pair of major salivary glands located inferior and anterior to 
the ears 


permanent tooth 
one of 32 adult teeth 


pharynx 
throat 


premolar 
(also, bicuspid) transitional tooth used for mastication, crushing, and 
grinding food 


pulp cavity 
deepest portion of a tooth, containing nerve endings and blood vessels 


root 
portion of a tooth embedded in the alveolar processes beneath the gum 
line 


saliva 
aqueous solution of proteins and ions secreted into the mouth by the 
salivary glands 


salivary amylase 
digestive enzyme in saliva that acts on starch 


salivary gland 
an exocrine gland that secretes a digestive fluid called saliva 


salivation 
secretion of saliva 


soft palate 
posterior region of the bottom portion of the nasal cavity that consists 
of skeletal muscle 


sublingual gland 
one of a pair of major salivary glands located beneath the tongue 


submandibular gland 
one of a pair of major salivary glands located in the floor of the mouth 


tongue 
accessory digestive organ of the mouth, the bulk of which is composed 
of skeletal muscle 


upper esophageal sphincter 
skeletal muscle sphincter that regulates food movement from the 
pharynx to the esophagus 


voluntary phase 
initial phase of deglutition, in which the bolus moves from the mouth 
to the oropharynx 


The Stomach 
By the end of this section, you will be able to: 


e Label on a diagram the four main regions of the stomach, its curvatures, and its 
sphincter 

e Identify the four main types of secreting cells in gastric glands, and their important 
products 

e Explain why the stomach does not digest itself 

¢ Describe the mechanical and chemical digestion of food entering the stomach 


Although a minimal amount of carbohydrate digestion occurs in the mouth, chemical 
digestion really gets underway in the stomach. An expansion of the alimentary canal that 
lies immediately inferior to the esophagus, the stomach links the esophagus to the first 
part of the small intestine (the duodenum) and is relatively fixed in place at its esophageal 
and duodenal ends. In between, however, it can be a highly active structure, contracting 
and continually changing position and size. These contractions provide mechanical 
assistance to digestion. The empty stomach is only about the size of your fist, but can 
stretch to hold as much as 4 liters of food and fluid, or more than 75 times its empty 
volume, and then return to its resting size when empty. Although you might think that the 
size of a person’s stomach is related to how much food that individual consumes, body 
weight does not correlate with stomach size. Rather, when you eat greater quantities of 
food—such as at holiday dinner—you stretch the stomach more than when you eat less. 


Popular culture tends to refer to the stomach as the location where all digestion takes 
place. Of course, this is not true. An important function of the stomach is to serve as a 
temporary holding chamber. You can ingest a meal far more quickly than it can be 
digested and absorbed by the small intestine. Thus, the stomach holds food and parses 
only small amounts into the small intestine at a time. Foods are not processed in the order 
they are eaten; rather, they are mixed together with digestive juices in the stomach until 
they are converted into chyme, which is released into the small intestine. 


As you will see in the sections that follow, the stomach plays several important roles in 
chemical digestion, including the continued digestion of carbohydrates and the initial 
digestion of proteins and triglycerides. Little if any nutrient absorption occurs in the 
stomach, with the exception of the negligible amount of nutrients in alcohol. 


Structure 


There are four main regions in the stomach: the cardia, fundus, body, and pylorus 
([link]). The cardia (or cardiac region) is the point where the esophagus connects to the 
stomach and through which food passes into the stomach. Located inferior to the 
diaphragm, above and to the left of the cardia, is the dome-shaped fundus. Below the 
fundus is the body, the main part of the stomach. The funnel-shaped pylorus connects 
the stomach to the duodenum. The wider end of the funnel, the pyloric antrum, connects 


to the body of the stomach. The narrower end is called the pyloric canal, which connects 
to the duodenum. The smooth muscle pyloric sphincter is located at this latter point of 
connection and controls stomach emptying. In the absence of food, the stomach deflates 
inward, and its mucosa and submucosa fall into a large fold called a ruga. 


Stomach 
Cardia 


Esophagus 


Muscularis externa: 
Longitudinal layer 
Circular layer 
Oblique layer 


Fundus 


Serosa 


Lesser curvature Body 


Pyloric sphincter 


(valve) at pylorus Lumen 


Rugae of mucosa 


Duodenum F ~~ 
Pyloric canal 


Pyloric antrum Greater curvature 


The stomach has four major regions: the cardia, fundus, 
body, and pylorus. The addition of an inner oblique smooth 
muscle layer gives the muscularis the ability to vigorously 

churn and mix food. 


The convex lateral surface of the stomach is called the greater curvature; the concave 
medial border is the lesser curvature. The stomach is held in place by the lesser omentum, 
which extends from the liver to the lesser curvature, and the greater omentum, which runs 
from the greater curvature to the posterior abdominal wall. 


Histology 


The wall of the stomach is made of the same four layers as most of the rest of the 
alimentary canal, but with adaptations to the mucosa and muscularis for the unique 
functions of this organ. In addition to the typical circular and longitudinal smooth muscle 
layers, the muscularis has an inner oblique smooth muscle layer ({link]). As a result, in 
addition to moving food through the canal, the stomach can vigorously churn food, 
mechanically breaking it down into smaller particles. 

Histology of the Stomach 


Parietal cell 
Surface 
epithelium 


Gastric pit 


Gastric gland 


Lamina 
propria 


Chief cell 


Muscularis 
mucosae 


Submucosa ———_—— aS > 
Oblique layer . 
Muscularis Circular layer —9 : » | Enteroendocrine 
externa Longitudinal SS / cell 
layer ~ y 


Serosa 


The stomach wall is adapted for the functions of the 
stomach. In the epithelium, gastric pits lead to gastric 
glands that secrete gastric juice. The gastric glands (one 
gland is shown enlarged on the right) contain different 
types of cells that secrete a variety of enzymes, including 
hydrochloride acid, which activates the protein-digesting 
enzyme pepsin. 


The stomach mucosa’s epithelial lining consists only of surface mucus cells, which 
secrete a protective coat of alkaline mucus. A vast number of gastric pits dot the surface 
of the epithelium, giving it the appearance of a well-used pincushion, and mark the entry 
to each gastric gland, which secretes a complex digestive fluid referred to as gastric 
juice. 


Although the walls of the gastric pits are made up primarily of mucus cells, the gastric 
glands are made up of different types of cells. The glands of the cardia and pylorus are 
composed primarily of mucus-secreting cells. Cells that make up the pyloric antrum 
secrete mucus and a number of hormones, including the majority of the stimulatory 
hormone, gastrin. The much larger glands of the fundus and body of the stomach, the site 
of most chemical digestion, produce most of the gastric secretions. These glands are 
made up of a variety of secretory cells. These include parietal cells, chief cells, mucous 
neck cells, and enteroendocrine cells. 


Parietal cells—Located primarily in the middle region of the gastric glands are parietal 
cells, which are among the most highly differentiated of the body’s epithelial cells. These 
relatively large cells produce both hydrochloric acid (HCI) and intrinsic factor. HCl is 
responsible for the high acidity (pH 1.5 to 3.5) of the stomach contents and is needed to 
activate the protein-digesting enzyme, pepsin. The acidity also kills much of the bacteria 
you ingest with food and helps to denature proteins, making them more available for 
enzymatic digestion. Intrinsic factor is a glycoprotein necessary for the absorption of 
vitamin Bj, in the small intestine. 


Chief cells—Located primarily in the basal regions of gastric glands are chief cells, 
which secrete pepsinogen, the inactive proenzyme form of pepsin. HCl is necessary for 
the conversion of pepsinogen to pepsin. 


Mucous neck cells—Gastric glands in the upper part of the stomach contain mucous 
neck cells that secrete thin, acidic mucus that is much different from the mucus secreted 
by the goblet cells of the surface epithelium. The role of this mucus is not currently 
known. 


Enteroendocrine cells—Finally, enteroendocrine cells found in the gastric glands secrete 
various hormones into the interstitial fluid of the lamina propria. These include gastrin, 
which is released mainly by enteroendocrine G cells. 


[link] describes the digestive functions of important hormones secreted by the stomach. 


Note: 


Op eC 
=> openstax COLLEGE 
arene: 

Watch this animation that depicts the structure of the stomach and how this structure 


functions in the initiation of protein digestion. This view of the stomach shows the 
characteristic rugae. What is the function of these rugae? 


Hormones Secreted by the Stomach 


Production Production 
Hormone site stimulus Target organ Action 


Hormones Secreted by the Stomach 


Hormone 


Gastrin 


Gastrin 


Gastrin 


Gastrin 


Ghrelin 


Production 


site 


Stomach 
mucosa, 
mainly G 
cells of the 
pyloric 
antrum 


Stomach 
mucosa, 
mainly G 
cells of the 
pyloric 
antrum 


Stomach 
mucosa, 
mainly G 
cells of the 
pyloric 
antrum 


Stomach 
mucosa, 
mainly G 
cells of the 
pyloric 
antrum 


Stomach 
mucosa, 
mainly 
fundus 


Production 
stimulus 


Presence of 
peptides 
and amino 
acids in 
stomach 


Presence of 
peptides 
and amino 
acids in 
stomach 


Presence of 
peptides 
and amino 
acids in 
stomach 


Presence of 
peptides 
and amino 
acids in 
stomach 


Fasting 
state (levels 
increase 
just prior to 
meals) 


Target organ 


Stomach 


Small 
intestine 


Ileocecal 
valve 


Large 
intestine 


Hypothalamus 


Action 


Increases 
secretion 
by gastric 
glands; 

promotes 
gastric 

emptying 


Promotes 
intestinal 
muscle 
contraction 


Relaxes 
valve 


Triggers 
mass 
movements 


Regulates 
food 
intake, 
primarily 
by 
stimulating 
hunger and 
satiety 


Hormones Secreted by the Stomach 


Hormone 


Histamine 


Serotonin 


Somatostatin 


Somatostatin 


Somatostatin 


Gastric Secretion 


Production 


site 


Stomach 
mucosa 


Stomach 
mucosa 


Mucosa of 
stomach, 
especially 
pyloric 
antrum; 
also 
duodenum 


Mucosa of 
stomach, 
especially 
pyloric 
antrum; 
also 
duodenum 


Mucosa of 
stomach, 
especially 
pyloric 
antrum; 
also 
duodenum 


Production 
stimulus 


Presence of 
food in the 
stomach 


Presence of 
food in the 
stomach 


Presence of 
food in the 
stomach; 
sympathetic 
axon 
stimulation 


Presence of 
food in the 
stomach; 
sympathetic 
axon 
stimulation 


Presence of 
food in the 
stomach; 
sympathetic 
axon 
stimulation 


Target organ 


Stomach 


Stomach 


Stomach 


Pancreas 


Small 
intestine 


Action 


Stimulates 
parietal 
cells to 
release 
HCl 


Contracts 
stomach 
muscle 


Restricts 
all gastric 
secretions, 
gastric 
motility, 
and 
emptying 


Restricts 
pancreatic 
secretions 


Reduces 
intestinal 
absorption 
by 
reducing 
blood flow 


The secretion of gastric juice is controlled by both nerves and hormones. Stimuli in the 
brain, stomach, and small intestine activate or inhibit gastric juice production. This is 
why the three phases of gastric secretion are called the cephalic, gastric, and intestinal 
phases ([link]). However, once gastric secretion begins, all three phases can occur 
simultaneously. 

The Three Phases of Gastric Secretion 


Stimulates stomach 
secretory activity 


CEPHALIC PHASE 


Inhibits stomach 
secretory activity 


Stimulates stomach 
secretory activity 


GASTRIC PHASE 


Inhibits stomach 
secretory activity 


Stimulates stomach 
secretory activity 


INTESTINAL PHASE 


Inhibits stomach 
secretory activity 


Gastric secretion occurs in three phases: cephalic, 
gastric, and intestinal. During each phase, the 
secretion of gastric juice can be stimulated or 

inhibited. 


The cephalic phase (reflex phase) of gastric secretion, which is relatively brief, takes 
place before food enters the stomach. The smell, taste, sight, or thought of food triggers 


this phase. For example, when you bring a piece of sushi to your lips, impulses from 
receptors in your taste buds or the nose are relayed to your brain, which returns signals 
that increase gastric secretion to prepare your stomach for digestion. This enhanced 
secretion is a conditioned reflex, meaning it occurs only if you like or want a particular 
food. Depression and loss of appetite can suppress the cephalic reflex. 


The gastric phase of secretion lasts 3 to 4 hours, and is set in motion by local neural and 
hormonal mechanisms triggered by the entry of food into the stomach. For example, 
when your sushi reaches the stomach, it creates distention that activates the stretch 
receptors. This stimulates parasympathetic neurons to release acetylcholine, which then 
provokes increased secretion of gastric juice. Partially digested proteins, caffeine, and 
rising pH stimulate the release of gastrin from enteroendocrine G cells, which in turn 
induces parietal cells to increase their production of HCl, which is needed to create an 
acidic environment for the conversion of pepsinogen to pepsin, and protein digestion. 
Additionally, the release of gastrin activates vigorous smooth muscle contractions. 
However, it should be noted that the stomach does have a natural means of avoiding 
excessive acid secretion and potential heartburn. Whenever pH levels drop too low, cells 
in the stomach react by suspending HCI secretion and increasing mucous secretions. 


The intestinal phase of gastric secretion has both excitatory and inhibitory elements. The 
duodenum has a major role in regulating the stomach and its emptying. When partially 
digested food fills the duodenum, intestinal mucosal cells release a hormone called 
intestinal (enteric) gastrin, which further excites gastric juice secretion. This stimulatory 
activity is brief, however, because when the intestine distends with chyme, the 
enterogastric reflex inhibits secretion. One of the effects of this reflex is to close the 
pyloric sphincter, which blocks additional chyme from entering the duodenum. 


The Mucosal Barrier 


The mucosa of the stomach is exposed to the highly corrosive acidity of gastric juice. 
Gastric enzymes that can digest protein can also digest the stomach itself. The stomach is 
protected from self-digestion by the mucosal barrier. This barrier has several 
components. First, the stomach wall is covered by a thick coating of bicarbonate-rich 
mucus. This mucus forms a physical barrier, and its bicarbonate ions neutralize acid. 
Second, the epithelial cells of the stomach's mucosa meet at tight junctions, which block 
gastric juice from penetrating the underlying tissue layers. Finally, stem cells located 
where gastric glands join the gastric pits quickly replace damaged epithelial mucosal 
cells, when the epithelial cells are shed. In fact, the surface epithelium of the stomach is 
completely replaced every 3 to 6 days. 


Note: 
Homeostatic Imbalances 


Ulcers: When the Mucosal Barrier Breaks Down 

As effective as the mucosal barrier is, it is not a “fail-safe” mechanism. Sometimes, 
gastric juice eats away at the superficial lining of the stomach mucosa, creating erosions, 
which mostly heal on their own. Deeper and larger erosions are called ulcers. 

Why does the mucosal barrier break down? A number of factors can interfere with its 
ability to protect the stomach lining. The majority of all ulcers are caused by either 
excessive intake of non-steroidal anti-inflammatory drugs (NSAIDs), including aspirin, 
or Helicobacter pylori infection. 

Antacids help relieve symptoms of ulcers such as “burning” pain and indigestion. When 
ulcers are caused by NSAID use, switching to other classes of pain relievers allows 
healing. When caused by H. pylori infection, antibiotics are effective. 

A potential complication of ulcers is perforation: Perforated ulcers create a hole in the 
stomach wall, resulting in peritonitis (inflammation of the peritoneum). These ulcers 
must be repaired surgically. 


Digestive Functions of the Stomach 


The stomach participates in virtually all the digestive activities with the exception of 
ingestion and defecation. Although almost all absorption takes place in the small 
intestine, the stomach does absorb some nonpolar substances, such as alcohol and aspirin. 


Mechanical Digestion 


Within a few moments after food after enters your stomach, mixing waves begin to occur 
at intervals of approximately 20 seconds. A mixing wave is a unique type of peristalsis 
that mixes and softens the food with gastric juices to create chyme. The initial mixing 
waves are relatively gentle, but these are followed by more intense waves, starting at the 
body of the stomach and increasing in force as they reach the pylorus. It is fair to say that 
long before your sushi exits through the pyloric sphincter, it bears little resemblance to 
the sushi you ate. 


The pylorus, which holds around 30 mL (1 fluid ounce) of chyme, acts as a filter, 
permitting only liquids and small food particles to pass through the mostly, but not fully, 
closed pyloric sphincter. In a process called gastric emptying, rhythmic mixing waves 
force about 3 mL of chyme at a time through the pyloric sphincter and into the 
duodenum. Release of a greater amount of chyme at one time would overwhelm the 
capacity of the small intestine to handle it. The rest of the chyme is pushed back into the 
body of the stomach, where it continues mixing. This process is repeated when the next 
mixing waves force more chyme into the duodenum. 


Gastric emptying is regulated by both the stomach and the duodenum. The presence of 
chyme in the duodenum activates receptors that inhibit gastric secretion. This prevents 
additional chyme from being released by the stomach before the duodenum is ready to 
process it. 


Chemical Digestion 


The fundus plays an important role, because it stores both undigested food and gases that 
are released during the process of chemical digestion. Food may sit in the fundus of the 
stomach for a while before being mixed with the chyme. While the food is in the fundus, 
the digestive activities of salivary amylase continue until the food begins mixing with the 
acidic chyme. Ultimately, mixing waves incorporate this food with the chyme, the acidity 
of which inactivates salivary amylase and activates lingual lipase. Lingual lipase then 
begins breaking down triglycerides into free fatty acids, and mono- and diglycerides. 


The breakdown of protein begins in the stomach through the actions of HCl and the 
enzyme pepsin. During infancy, gastric glands also produce rennin, an enzyme that helps 
digest milk protein. 


Its numerous digestive functions notwithstanding, there is only one stomach function 
necessary to life: the production of intrinsic factor. The intestinal absorption of vitamin 
By», which is necessary for both the production of mature red blood cells and normal 
neurological functioning, cannot occur without intrinsic factor. People who undergo total 
gastrectomy (stomach removal)—for life-threatening stomach cancer, for example—can 
survive with minimal digestive dysfunction if they receive vitamin B,> injections. 


The contents of the stomach are completely emptied into the duodenum within 2 to 4 
hours after you eat a meal. Different types of food take different amounts of time to 
process. Foods heavy in carbohydrates empty fastest, followed by high-protein foods. 
Meals with a high triglyceride content remain in the stomach the longest. Since enzymes 
in the small intestine digest fats slowly, food can stay in the stomach for 6 hours or longer 
when the duodenum is processing fatty chyme. However, note that this is still a fraction 
of the 24 to 72 hours that full digestion typically takes from start to finish. 


Chapter Review 


The stomach participates in all digestive activities except ingestion and defecation. It 
vigorously churns food. It secretes gastric juices that break down food and absorbs 
certain drugs, including aspirin and some alcohol. The stomach begins the digestion of 
protein and continues the digestion of carbohydrates and fats. It stores food as an acidic 
liquid called chyme, and releases it gradually into the small intestine through the pyloric 
sphincter. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this animation that depicts the structure of the stomach and how this structure 
functions in the initiation of protein digestion. This view of the stomach shows the 
characteristic rugae. What is the function of these rugae? 


Solution: 


Answers may vary. 


Review Questions 


Exercise: 


Problem: Which of these cells secrete hormones? 


a. parietal cells 

b. mucous neck cells 

c. enteroendocrine cells 
d. chief cells 


Solution: 


C 


Exercise: 


Problem:Where does the majority of chemical digestion in the stomach occur? 


a. fundus and body 
b. cardia and fundus 
c. body and pylorus 
d. body 


Solution: 


A 


Exercise: 


Problem: 


During gastric emptying, chyme is released into the duodenum through the 


a. esophageal hiatus 
b. pyloric antrum 

c. pyloric canal 

d. pyloric sphincter 


Solution: 


D 


Exercise: 


Problem: Parietal cells secrete 


a. gastrin 

b. hydrochloric acid 
c. pepsin 

d. pepsinogen 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


Explain how the stomach is protected from self-digestion and why this is necessary. 
Solution: 


The mucosal barrier protects the stomach from self-digestion. It includes a thick 
coating of bicarbonate-rich mucus; the mucus is physically protective, and 
bicarbonate neutralizes gastric acid. Epithelial cells meet at tight junctions, which 
block gastric juice from penetrating the underlying tissue layers, and stem cells 
quickly replace sloughed off epithelial mucosal cells. 


Exercise: 


Problem: 


Describe unique anatomical features that enable the stomach to perform digestive 
functions. 


Solution: 


The stomach has an additional inner oblique smooth muscle layer that helps the 
muscularis churn and mix food. The epithelium includes gastric glands that secrete 
gastric fluid. The gastric fluid consists mainly of mucous, HCl, and the enzyme 
pepsin released as pepsinogen. 


Glossary 


body 
mid-portion of the stomach 


cardia 
(also, cardiac region) part of the stomach surrounding the cardiac orifice (esophageal 
hiatus) 


cephalic phase 
(also, reflex phase) initial phase of gastric secretion that occurs before food enters 
the stomach 


chief cell 
gastric gland cell that secretes pepsinogen 


enteroendocrine cell 
gastric gland cell that releases hormones 


fundus 
dome-shaped region of the stomach above and to the left of the cardia 


G cell 
gastrin-secreting enteroendocrine cell 


gastric emptying 
process by which mixing waves gradually cause the release of chyme into the 
duodenum 


gastric gland 
gland in the stomach mucosal epithelium that produces gastric juice 


gastric phase 
phase of gastric secretion that begins when food enters the stomach 


gastric pit 
narrow channel formed by the epithelial lining of the stomach mucosa 


gastrin 
peptide hormone that stimulates secretion of hydrochloric acid and gut motility 


hydrochloric acid (HCl) 
digestive acid secreted by parietal cells in the stomach 


intrinsic factor 
glycoprotein required for vitamin B,> absorption in the small intestine 


intestinal phase 
phase of gastric secretion that begins when chyme enters the intestine 


mixing wave 
unique type of peristalsis that occurs in the stomach 


mucosal barrier 
protective barrier that prevents gastric juice from destroying the stomach itself 


mucous neck cell 
gastric gland cell that secretes a uniquely acidic mucus 


parietal cell 
gastric gland cell that secretes hydrochloric acid and intrinsic factor 


pepsinogen 
inactive form of pepsin 


pyloric antrum 
wider, more superior part of the pylorus 


pyloric canal 
narrow, more inferior part of the pylorus 


pyloric sphincter 
sphincter that controls stomach emptying 


pylorus 
lower, funnel-shaped part of the stomach that is continuous with the duodenum 


ruga 


fold of alimentary canal mucosa and submucosa in the empty stomach and other 
organs 


stomach 
alimentary canal organ that contributes to chemical and mechanical digestion of 
food from the esophagus before releasing it, as chyme, to the small intestine 


The Small and Large Intestines 
By the end of this section, you will be able to: 


¢ Compare and contrast the location and gross anatomy of the small and 
large intestines 

e Identify three main adaptations of the small intestine wall that increase 
its absorptive capacity 

e Describe the mechanical and chemical digestion of chyme upon its 
release into the small intestine 

e List three features unique to the wall of the large intestine and identify 
their contributions to its function 

e Identify the beneficial roles of the bacterial flora in digestive system 
functioning 

e Trace the pathway of food waste from its point of entry into the large 
intestine through its exit from the body as feces 


The word intestine is derived from a Latin root meaning “internal,” and 
indeed, the two organs together nearly fill the interior of the abdominal 
cavity. In addition, called the small and large bowel, or colloquially the 
“suts,” they constitute the greatest mass and length of the alimentary canal 
and, with the exception of ingestion, perform all digestive system functions. 


The Small Intestine 


Chyme released from the stomach enters the small intestine, which is the 
primary digestive organ in the body. Not only is this where most digestion 
occurs, it is also where practically all absorption occurs. The longest part of 
the alimentary canal, the small intestine is about 3.05 meters (10 feet) long 
in a living person (but about twice as long in a cadaver due to the loss of 
muscle tone). Since this makes it about five times longer than the large 
intestine, you might wonder why it is called “small.” In fact, its name 
derives from its relatively smaller diameter of only about 2.54 cm (1 in), 
compared with 7.62 cm (3 in) for the large intestine. As we’ll see shortly, in 
addition to its length, the folds and projections of the lining of the small 
intestine work to give it an enormous surface area, which is approximately 
200 m?, more than 100 times the surface area of your skin. This large 


surface area is necessary for complex processes of digestion and absorption 
that occur within it. 


Structure 


The coiled tube of the small intestine is subdivided into three regions. From 
proximal (at the stomach) to distal, these are the duodenum, jejunum, and 
ileum ([link]). 


The shortest region is the 25.4-cm (10-in) duodenum, which begins at the 
pyloric sphincter. Just past the pyloric sphincter, it bends posteriorly behind 
the peritoneum, becoming retroperitoneal, and then makes a C-shaped curve 
around the head of the pancreas before ascending anteriorly again to return 
to the peritoneal cavity and join the jejunum. The duodenum can therefore 
be subdivided into four segments: the superior, descending, horizontal, and 
ascending duodenum. 


Of particular interest is the hepatopancreatic ampulla (ampulla of Vater). 
Located in the duodenal wall, the ampulla marks the transition from the 
anterior portion of the alimentary canal to the mid-region, and is where the 
bile duct (through which bile passes from the liver) and the main 
pancreatic duct (through which pancreatic juice passes from the pancreas) 
join. This ampulla opens into the duodenum at a tiny volcano-shaped 
structure called the major duodenal papilla. The hepatopancreatic 
sphincter (sphincter of Oddi) regulates the flow of both bile and pancreatic 
juice from the ampulla into the duodenum. 

Small Intestine 


Duodenum 
Jejunum 
lleum 
Large intestine 


Rectum ———___—>=_ 


The three regions of the small intestine are the 
duodenum, jejunum, and ileum. 


The jejunum is about 0.9 meters (3 feet) long (in life) and runs from the 
duodenum to the ileum. Jejunum means “empty” in Latin and supposedly 
was so named by the ancient Greeks who noticed it was always empty at 
death. No clear demarcation exists between the jejunum and the final 
segment of the small intestine, the ileum. 


The ileum is the longest part of the small intestine, measuring about 1.8 
meters (6 feet) in length. It is thicker, more vascular, and has more 
developed mucosal folds than the jejunum. The ileum joins the cecum, the 
first portion of the large intestine, at the ileocecal sphincter (or valve). The 
jejunum and ileum are tethered to the posterior abdominal wall by the 


mesentery. The large intestine frames these three parts of the small 
intestine. 


Parasympathetic nerve fibers from the vagus nerve and sympathetic nerve 
fibers from the thoracic splanchnic nerve provide extrinsic innervation to 
the small intestine. The superior mesenteric artery is its main arterial 
supply. Veins run parallel to the arteries and drain into the superior 


mesenteric vein. Nutrient-rich blood from the small intestine is then carried 
to the liver via the hepatic portal vein. 


Histology 


The wall of the small intestine is composed of the same four layers typically 
present in the alimentary system. However, three features of the mucosa and 
submucosa are unique. These features, which increase the absorptive 
surface area of the small intestine more than 600-fold, include circular 
folds, villi, and microvilli ([link]). These adaptations are most abundant in 
the proximal two-thirds of the small intestine, where the majority of 
absorption occurs. 

~~ of the Small ie 


Absorptive cells 


Capillary oe 
Artery (brush border) 


Goblet cell 


trl@t JURee Jet Be! 


° ony OS ED Vac 


|i 


Lymphatic vesicle 


Muscularis mucosae Duodenal gland 


(a) 


(a) The absorptive surface of the small intestine is vastly 
enlarged by the presence of circular folds, villi, and microvilli. 
(b) Micrograph of the circular folds. (c) Micrograph of the villi. 


(d) Electron micrograph of the microvilli. From left to right, 

LM x 56, LM x 508, EM x 196,000. (credit b-d: Micrograph 

provided by the Regents of University of Michigan Medical 
School © 2012) 


Circular folds 


Also called a plica circulare, a circular fold is a deep ridge in the mucosa 
and submucosa. Beginning near the proximal part of the duodenum and 
ending near the middle of the ileum, these folds facilitate absorption. Their 
shape causes the chyme to spiral, rather than move in a straight line, 
through the small intestine. Spiraling slows the movement of chyme and 
provides the time needed for nutrients to be fully absorbed. 


Villi 


Within the circular folds are small (0.5—1 mm long) hairlike vascularized 
projections called villi (singular = villus) that give the mucosa a furry 
texture. There are about 20 to 40 villi per square millimeter, increasing the 
surface area of the epithelium tremendously. The mucosal epithelium, 
primarily composed of absorptive cells, covers the villi. In addition to 
muscle and connective tissue to support its structure, each villus contains a 
capillary bed composed of one arteriole and one venule, as well as a 
lymphatic capillary called a lacteal. The breakdown products of 
carbohydrates and proteins (sugars and amino acids) can enter the 
bloodstream directly, but lipid breakdown products are absorbed by the 
lacteals and transported to the bloodstream via the lymphatic system. 


Microvilli 


As their name suggests, microvilli (singular = microvillus) are much 
smaller (1 pm) than villi. They are cylindrical apical surface extensions of 
the plasma membrane of the mucosa’s epithelial cells, and are supported by 
microfilaments within those cells. Although their small size makes it 
difficult to see each microvillus, their combined microscopic appearance 
suggests a mass of bristles, which is termed the brush border. Fixed to the 
surface of the microvilli membranes are enzymes that finish digesting 
carbohydrates and proteins. There are an estimated 200 million microvilli 
per square millimeter of small intestine, greatly expanding the surface area 
of the plasma membrane and thus greatly enhancing absorption. 


Intestinal Glands 


In addition to the three specialized absorptive features just discussed, the 
mucosa between the villi is dotted with deep crevices that each lead into a 
tubular intestinal gland (crypt of Lieberkiihn), which is formed by cells 
that line the crevices (see [link]). These produce intestinal juice, a slightly 
alkaline (pH 7.4 to 7.8) mixture of water and mucus. Each day, about 0.95 
to 1.9 liters (1 to 2 quarts) are secreted in response to the distention of the 
small intestine or the irritating effects of chyme on the intestinal mucosa. 


The submucosa of the duodenum is the only site of the complex mucus- 
secreting duodenal glands (Brunner’s glands), which produce a 
bicarbonate-rich alkaline mucus that buffers the acidic chyme as it enters 
from the stomach. 


The roles of the cells in the small intestinal mucosa are detailed in [link]. 


Cells of the Small Intestinal Mucosa 


Cells of the SmdlbhatéstialtNéucosa 


Cell type 


Cell type 


Absorptive 


Goblet 


Paneth 


G cells 


I cells 


K cells 


mucosa 


Location in the 
mucosa 


Epithelium/intestinal 
glands 


Epithelium/intestinal 
glands 


Intestinal glands 


Intestinal glands of 
duodenum 


Intestinal glands of 
duodenum 


Intestinal glands 


Function 


Function 


Digestion and absorption 
of nutrients in chyme 


Secretion of mucus 


Secretion of the 
bactericidal enzyme 
lysozyme; phagocytosis 


Secretion of the hormone 
intestinal gastrin 


Secretion of the hormone 
cholecystokinin, which 
stimulates release of 
pancreatic juices and bile 


Secretion of the hormone 
glucose-dependent 
insulinotropic peptide, 
which stimulates the 
release of insulin 


Cells of the Small Intestinal Mucosa 


Location in the 
Cell type mucosa Function 


Secretion of the hormone 
motilin, which accelerates 


Intestinal glands of ; 
gastric emptying, 


M cells duodenum and : ; ‘ 
oe stimulates intestinal 
jejunum ; : : 
peristalsis, and stimulates 
the production of pepsin 
? retion of the h 
S cells Intestinal glands Sec eno Die OENOn’ 
secretin 
Intestinal MALT 


The lamina propria of the small intestine mucosa is studded with quite a bit 
of MALT. In addition to solitary lymphatic nodules, aggregations of 
intestinal MALT, which are typically referred to as Peyer’s patches, are 
concentrated in the distal ileum, and serve to keep bacteria from entering 
the bloodstream. Peyer’s patches are most prominent in young people and 
become less distinct as you age, which coincides with the general activity of 
our immune system. 


Note: 


— 
meee OPENStAX COLLEGE 


Watch this animation that depicts the structure of the small intestine, and, 
in particular, the villi. Epithelial cells continue the digestion and absorption 
of nutrients and transport these nutrients to the lymphatic and circulatory 
systems. In the small intestine, the products of food digestion are absorbed 
by different structures in the villi. Which structure absorbs and transports 
fats? 


Mechanical Digestion in the Small Intestine 


The movement of intestinal smooth muscles includes both segmentation 
and a form of peristalsis called migrating motility complexes. The kind of 
peristaltic mixing waves seen in the stomach are not observed here. 


If you could see into the small intestine when it was going through 
segmentation, it would look as if the contents were being shoved 
incrementally back and forth, as the rings of smooth muscle repeatedly 
contract and then relax. Segmentation in the small intestine does not force 
chyme through the tract. Instead, it combines the chyme with digestive 
juices and pushes food particles against the mucosa to be absorbed. The 
duodenum is where the most rapid segmentation occurs, at a rate of about 
12 times per minute. In the ileum, segmentations are only about eight times 
per minute ([link]). 

Segmentation 


Segmentation separates 
chyme and then pushes it 
back together, mixing it 
and providing time for 
digestion and absorption. 


When most of the chyme has been absorbed, the small intestinal wall 
becomes less distended. At this point, the localized segmentation process is 
replaced by transport movements. The duodenal mucosa secretes the 
hormone motilin, which initiates peristalsis in the form of a migrating 
motility complex. These complexes, which begin in the duodenum, force 
chyme through a short section of the small intestine and then stop. The next 
contraction begins a little bit farther down than the first, forces chyme a bit 
farther through the small intestine, then stops. These complexes move 
slowly down the small intestine, forcing chyme on the way, taking around 
90 to 120 minutes to finally reach the end of the ileum. At this point, the 
process is repeated, starting in the duodenum. 


The ileocecal valve, a sphincter, is usually in a constricted state, but when 
motility in the ileum increases, this sphincter relaxes, allowing food residue 
to enter the first portion of the large intestine, the cecum. Relaxation of the 
ileocecal sphincter is controlled by both nerves and hormones. First, 


digestive activity in the stomach provokes the gastroileal reflex, which 
increases the force of ileal segmentation. Second, the stomach releases the 
hormone gastrin, which enhances ileal motility, thus relaxing the ileocecal 
sphincter. After chyme passes through, backward pressure helps close the 
sphincter, preventing backflow into the ileum. Because of this reflex, your 
lunch is completely emptied from your stomach and small intestine by the 
time you eat your dinner. It takes about 3 to 5 hours for all chyme to leave 
the small intestine. 


Chemical Digestion in the Small Intestine 


The digestion of proteins and carbohydrates, which partially occurs in the 
stomach, is completed in the small intestine with the aid of intestinal and 
pancreatic juices. Lipids arrive in the intestine largely undigested, so much 
of the focus here is on lipid digestion, which is facilitated by bile and the 
enzyme pancreatic lipase. 


Moreover, intestinal juice combines with pancreatic juice to provide a liquid 
medium that facilitates absorption. The intestine is also where most water is 
absorbed, via osmosis. The small intestine’s absorptive cells also synthesize 
digestive enzymes and then place them in the plasma membranes of the 
microvilli. This distinguishes the small intestine from the stomach; that is, 
enzymatic digestion occurs not only in the lumen, but also on the luminal 
surfaces of the mucosal cells. 


For optimal chemical digestion, chyme must be delivered from the stomach 
slowly and in small amounts. This is because chyme from the stomach is 
typically hypertonic, and if large quantities were forced all at once into the 
small intestine, the resulting osmotic water loss from the blood into the 
intestinal lumen would result in potentially life-threatening low blood 
volume. In addition, continued digestion requires an upward adjustment of 
the low pH of stomach chyme, along with rigorous mixing of the chyme 
with bile and pancreatic juices. Both processes take time, so the pumping 
action of the pylorus must be carefully controlled to prevent the duodenum 
from being overwhelmed with chyme. 


Note: 

Disorders of the... 

Small Intestine: Lactose Intolerance 

Lactose intolerance is a condition characterized by indigestion caused by 
dairy products. It occurs when the absorptive cells of the small intestine do 
not produce enough lactase, the enzyme that digests the milk sugar lactose. 
In most mammals, lactose intolerance increases with age. In contrast, some 
human populations, most notably Caucasians, are able to maintain the 
ability to produce lactase as adults. 

In people with lactose intolerance, the lactose in chyme is not digested. 
Bacteria in the large intestine ferment the undigested lactose, a process that 
produces gas. In addition to gas, symptoms include abdominal cramps, 
bloating, and diarrhea. Symptom severity ranges from mild discomfort to 
severe pain; however, symptoms resolve once the lactose is eliminated in 
feces. 

The hydrogen breath test is used to help diagnose lactose intolerance. 
Lactose-tolerant people have very little hydrogen in their breath. Those 
with lactose intolerance exhale hydrogen, which is one of the gases 
produced by the bacterial fermentation of lactose in the colon. After the 
hydrogen is absorbed from the intestine, it is transported through blood 
vessels into the lungs. There are a number of lactose-free dairy products 
available in grocery stores. In addition, dietary supplements are available. 
Taken with food, they provide lactase to help digest lactose. 


The Large Intestine 


The large intestine is the terminal part of the alimentary canal. The primary 
function of this organ is to finish absorption of nutrients and water, 
synthesize certain vitamins, form feces, and eliminate feces from the body. 


Structure 


The large intestine runs from the appendix to the anus. It frames the small 
intestine on three sides. Despite its being about one-half as long as the small 


intestine, it is called large because it is more than twice the diameter of the 
small intestine, about 3 inches. 


Subdivisions 


The large intestine is subdivided into four main regions: the cecum, the 
colon, the rectum, and the anus. The ileocecal valve, located at the opening 
between the ileum and the large intestine, controls the flow of chyme from 
the small intestine to the large intestine. 


Cecum 


The first part of the large intestine is the cecum, a sac-like structure that is 
suspended inferior to the ileocecal valve. It is about 6 cm (2.4 in) long, 
receives the contents of the ileum, and continues the absorption of water 
and salts. The appendix (or vermiform appendix) is a winding tube that 
attaches to the cecum. Although the 7.6-cm (3-in) long appendix contains 
lymphoid tissue, suggesting an immunologic function, this organ is 
generally considered vestigial. However, at least one recent report 
postulates a survival advantage conferred by the appendix: In diarrheal 
illness, the appendix may serve as a bacterial reservoir to repopulate the 
enteric bacteria for those surviving the initial phases of the illness. 
Moreover, its twisted anatomy provides a haven for the accumulation and 
multiplication of enteric bacteria. The mesoappendix, the mesentery of the 
appendix, tethers it to the mesentery of the ileum. 


Colon 


The cecum blends seamlessly with the colon. Upon entering the colon, the 
food residue first travels up the ascending colon on the right side of the 
abdomen. At the inferior surface of the liver, the colon bends to form the 
right colic flexure (hepatic flexure) and becomes the transverse colon. 
The region defined as hindgut begins with the last third of the transverse 


colon and continues on. Food residue passing through the transverse colon 
travels across to the left side of the abdomen, where the colon angles 
sharply immediately inferior to the spleen, at the left colic flexure (splenic 
flexure). From there, food residue passes through the descending colon, 
which runs down the left side of the posterior abdominal wall. After 
entering the pelvis inferiorly, it becomes the s-shaped sigmoid colon, which 
extends medially to the midline ([{link]). The ascending and descending 
colon, and the rectum (discussed next) are located in the retroperitoneum. 
The transverse and sigmoid colon are tethered to the posterior abdominal 
wall by the mesocolon. 


Large Intestine 
Right colic 


(hepatic) flexure Left colic 
(splenic) 
flexure 

Transverse 

colon 

Ascending Descending 

colon colon 

lleum 

Cecum 

Vermiform Sigmoid 


appendix colon 


Anal canal Rectum 


The large intestine includes the 
cecum, colon, and rectum. 


Note: 

Homeostatic Imbalances 

Colorectal Cancer 

Each year, approximately 140,000 Americans are diagnosed with 
colorectal cancer, and another 49,000 die from it, making it one of the most 
deadly malignancies. People with a family history of colorectal cancer are 
at increased risk. Smoking, excessive alcohol consumption, and a diet high 
in animal fat and protein also increase the risk. Despite popular opinion to 


the contrary, studies support the conclusion that dietary fiber and calcium 
do not reduce the risk of colorectal cancer. 

Colorectal cancer may be signaled by constipation or diarrhea, cramping, 
abdominal pain, and rectal bleeding. Bleeding from the rectum may be 
either obvious or occult (hidden in feces). Since most colon cancers arise 
from benign mucosal growths called polyps, cancer prevention is focused 
on identifying these polyps. The colonoscopy is both diagnostic and 
therapeutic. Colonoscopy not only allows identification of precancerous 
polyps, the procedure also enables them to be removed before they become 
malignant. Screening for fecal occult blood tests and colonoscopy is 
recommended for those over 50 years of age. 


Rectum 


Food residue leaving the sigmoid colon enters the rectum in the pelvis, 
near the third sacral vertebra. The final 20.3 cm (8 in) of the alimentary 
canal, the rectum extends anterior to the sacrum and coccyx. Even though 
rectum is Latin for “straight,” this structure follows the curved contour of 
the sacrum and has three lateral bends that create a trio of internal 
transverse folds called the rectal valves. These valves help separate the 
feces from gas to prevent the simultaneous passage of feces and gas. 


Anal Canal 


Finally, food residue reaches the last part of the large intestine, the anal 
canal, which is located in the perineum, completely outside of the 
abdominopelvic cavity. This 3.8—5 cm (1.5—2 in) long structure opens to the 
exterior of the body at the anus. The anal canal includes two sphincters. The 
internal anal sphincter is made of smooth muscle, and its contractions are 
involuntary. The external anal sphincter is made of skeletal muscle, which 
is under voluntary control. Except when defecating, both usually remain 
closed. 


Histology 


There are several notable differences between the walls of the large and 
small intestines ([link]). For example, few enzyme-secreting cells are found 
in the wall of the large intestine, and there are no circular folds or villi. 
Other than in the anal canal, the mucosa of the colon is simple columnar 
epithelium made mostly of enterocytes (absorptive cells) and goblet cells. 
In addition, the wall of the large intestine has far more intestinal glands, 
which contain a vast population of enterocytes and goblet cells. These 
goblet cells secrete mucus that eases the movement of feces and protects the 
intestine from the effects of the acids and gases produced by enteric 
bacteria. The enterocytes absorb water and salts as well as vitamins 
produced by your intestinal bacteria. 

Histology of the large Intestine 


Openings of Microvilli 
intestinal glands ‘ond aN — = 
~~ NI \\(] Absorptive cell 


4 absorbs water 


Large intestine 


Goblet cell 
secretes mucus 


@We 
eas ] 
Smooth muscle fiber —————— ae 
Lymphatic nodule 


Muscularis mucosae 
Submucosa 


(a) The histologies of the large intestine and small 
intestine (not shown) are adapted for the digestive 
functions of each organ. (b) This micrograph shows 
the colon’s simple columnar epithelium and goblet 


cells. LM x 464. (credit b: Micrograph provided by the 
Regents of University of Michigan Medical School © 
2012) 


Anatomy 


Three features are unique to the large intestine: teniae coli, haustra, and 
epiploic appendages ([link]). The teniae coli are three bands of smooth 
muscle that make up the longitudinal muscle layer of the muscularis of the 
large intestine, except at its terminal end. Tonic contractions of the teniae 
coli bunch up the colon into a succession of pouches called haustra 
(singular = haustrum), which are responsible for the wrinkled appearance of 
the colon. Attached to the teniae coli are small, fat-filled sacs of visceral 
peritoneum called epiploic appendages. The purpose of these is unknown. 
Although the rectum and anal canal have neither teniae coli nor haustra, 
they do have well-developed layers of muscularis that create the strong 
contractions needed for defecation. 

Teniae Coli, Haustra, and Epiploic Appendages 


Epiploic appendages 


The stratified squamous epithelial mucosa of the anal canal connects to the 
skin on the outside of the anus. This mucosa varies considerably from that 
of the rest of the colon to accommodate the high level of abrasion as feces 
pass through. The anal canal’s mucous membrane is organized into 
longitudinal folds, each called an anal column, which house a grid of 
arteries and veins. Two superficial venous plexuses are found in the anal 
canal: one within the anal columns and one at the anus. 


Depressions between the anal columns, each called an anal sinus, secrete 
mucus that facilitates defecation. The pectinate line (or dentate line) is a 
horizontal, jagged band that runs circumferentially just below the level of 
the anal sinuses, and represents the junction between the hindgut and 
external skin. The mucosa above this line is fairly insensitive, whereas the 
area below is very sensitive. The resulting difference in pain threshold is 
due to the fact that the upper region is innervated by visceral sensory fibers, 
and the lower region is innervated by somatic sensory fibers. 


Bacterial Flora 


Most bacteria that enter the alimentary canal are killed by lysozyme, 
defensins, HCl, or protein-digesting enzymes. However, trillions of bacteria 
live within the large intestine and are referred to as the bacterial flora. 
Most of the more than 700 species of these bacteria are nonpathogenic 
commensal organisms that cause no harm as long as they stay in the gut 
lumen. In fact, many facilitate chemical digestion and absorption, and some 
synthesize certain vitamins, mainly biotin, pantothenic acid, and vitamin K. 
Some are linked to increased immune response. A refined system prevents 
these bacteria from crossing the mucosal barrier. First, peptidoglycan, a 
component of bacterial cell walls, activates the release of chemicals by the 
mucosa’s epithelial cells, which draft immune cells, especially dendritic 
cells, into the mucosa. Dendritic cells open the tight junctions between 
epithelial cells and extend probes into the lumen to evaluate the microbial 
antigens. The dendritic cells with antigens then travel to neighboring 
lymphoid follicles in the mucosa where T cells inspect for antigens. This 
process triggers an IgA-mediated response, if warranted, in the lumen that 


blocks the commensal organisms from infiltrating the mucosa and setting 
off a far greater, widespread systematic reaction. 


Digestive Functions of the Large Intestine 


The residue of chyme that enters the large intestine contains few nutrients 
except water, which is reabsorbed as the residue lingers in the large 
intestine, typically for 12 to 24 hours. Thus, it may not surprise you that the 
large intestine can be completely removed without significantly affecting 
digestive functioning. For example, in severe cases of inflammatory bowel 
disease, the large intestine can be removed by a procedure known as a 
colectomy. Often, a new fecal pouch can be crafted from the small intestine 
and sutured to the anus, but if not, an ileostomy can be created by bringing 
the distal ileum through the abdominal wall, allowing the watery chyme to 
be collected in a bag-like adhesive appliance. 


Mechanical Digestion 


In the large intestine, mechanical digestion begins when chyme moves from 
the ileum into the cecum, an activity regulated by the ileocecal sphincter. 
Right after you eat, peristalsis in the ileum forces chyme into the cecum. 
When the cecum is distended with chyme, contractions of the ileocecal 
sphincter strengthen. Once chyme enters the cecum, colon movements 
begin. 


Mechanical digestion in the large intestine includes a combination of three 
types of movements. The presence of food residues in the colon stimulates a 
slow-moving haustral contraction. This type of movement involves 
sluggish segmentation, primarily in the transverse and descending colons. 
When a haustrum is distended with chyme, its muscle contracts, pushing the 
residue into the next haustrum. These contractions occur about every 30 
minutes, and each last about 1 minute. These movements also mix the food 
residue, which helps the large intestine absorb water. The second type of 
movement is peristalsis, which, in the large intestine, is slower than in the 
more proximal portions of the alimentary canal. The third type is a mass 


movement. These strong waves start midway through the transverse colon 
and quickly force the contents toward the rectum. Mass movements usually 
occur three or four times per day, either while you eat or immediately 
afterward. Distension in the stomach and the breakdown products of 
digestion in the small intestine provoke the gastrocolic reflex, which 
increases motility, including mass movements, in the colon. Fiber in the diet 
both softens the stool and increases the power of colonic contractions, 
optimizing the activities of the colon. 


Chemical Digestion 


Although the glands of the large intestine secrete mucus, they do not secrete 
digestive enzymes. Therefore, chemical digestion in the large intestine 
occurs exclusively because of bacteria in the lumen of the colon. Through 
the process of saccharolytic fermentation, bacteria break down some of 
the remaining carbohydrates. This results in the discharge of hydrogen, 
carbon dioxide, and methane gases that create flatus (gas) in the colon; 
flatulence is excessive flatus. Each day, up to 1500 mL of flatus is produced 
in the colon. More is produced when you eat foods such as beans, which are 
rich in otherwise indigestible sugars and complex carbohydrates like 
soluble dietary fiber. 


Absorption, Feces Formation, and Defecation 


The small intestine absorbs about 90 percent of the water you ingest (either 
as liquid or within solid food). The large intestine absorbs most of the 
remaining water, a process that converts the liquid chyme residue into 
semisolid feces (“stool”). Feces is composed of undigested food residues, 
unabsorbed digested substances, millions of bacteria, old epithelial cells 
from the GI mucosa, inorganic salts, and enough water to let it pass 
smoothly out of the body. Of every 500 mL (17 ounces) of food residue that 
enters the cecum each day, about 150 mL (5 ounces) become feces. 


Feces are eliminated through contractions of the rectal muscles. You help 
this process by a voluntary procedure called Valsalva’s maneuver, in 


which you increase intra-abdominal pressure by contracting your 
diaphragm and abdominal wall muscles, and closing your glottis. 


The process of defecation begins when mass movements force feces from 
the colon into the rectum, stretching the rectal wall and provoking the 
defecation reflex, which eliminates feces from the rectum. This 
parasympathetic reflex is mediated by the spinal cord. It contracts the 
sigmoid colon and rectum, relaxes the internal anal sphincter, and initially 
contracts the external anal sphincter. The presence of feces in the anal canal 
sends a signal to the brain, which gives you the choice of voluntarily 
opening the external anal sphincter (defecating) or keeping it temporarily 
closed. If you decide to delay defecation, it takes a few seconds for the 
reflex contractions to stop and the rectal walls to relax. The next mass 
movement will trigger additional defecation reflexes until you defecate. 


If defecation is delayed for an extended time, additional water is absorbed, 
making the feces firmer and potentially leading to constipation. On the 
other hand, if the waste matter moves too quickly through the intestines, not 
enough water is absorbed, and diarrhea can result. This can be caused by 
the ingestion of foodborne pathogens. In general, diet, health, and stress 
determine the frequency of bowel movements. The number of bowel 
movements varies greatly between individuals, ranging from two or three 
per day to three or four per week. 


Note: 


eee 
—— 
mss" OPENStax COLLEGE 


By watching this animation you will see that for the various food groups— 
proteins, fats, and carbohydrates—digestion begins in different parts of the 
digestion system, though all end in the same place. Of the three major food 


classes (carbohydrates, fats, and proteins), which is digested in the mouth, 
the stomach, and the small intestine? 


Chapter Review 


The three main regions of the small intestine are the duodenum, the 
jejunum, and the ileum. The small intestine is where digestion is completed 
and virtually all absorption occurs. These two activities are facilitated by 
structural adaptations that increase the mucosal surface area by 600-fold, 
including circular folds, villi, and microvilli. There are around 200 million 
microvilli per square millimeter of small intestine, which contain brush 
border enzymes that complete the digestion of carbohydrates and proteins. 
Combined with pancreatic juice, intestinal juice provides the liquid medium 
needed to further digest and absorb substances from chyme. The small 
intestine is also the site of unique mechanical digestive movements. 
Segmentation moves the chyme back and forth, increasing mixing and 
opportunities for absorption. Migrating motility complexes propel the 
residual chyme toward the large intestine. 


The main regions of the large intestine are the cecum, the colon, and the 
rectum. The large intestine absorbs water and forms feces, and is 
responsible for defecation. Bacterial flora break down additional 
carbohydrate residue, and synthesize certain vitamins. The mucosa of the 
large intestinal wall is generously endowed with goblet cells, which secrete 
mucus that eases the passage of feces. The entry of feces into the rectum 
activates the defecation reflex. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this animation that depicts the structure of the small intestine, 
and, in particular, the villi. Epithelial cells continue the digestion and 
absorption of nutrients and transport these nutrients to the lymphatic 
and circulatory systems. In the small intestine, the products of food 
digestion are absorbed by different structures in the villi. Which 
structure absorbs and transports fats? 


Solution: 


Answers may vary. 

Exercise: 
Problem: 
By watching this animation, you will see that for the various food 
groups—proteins, fats, and carbohydrates—digestion begins in 
different parts of the digestion system, though all end in the same 
place. Of the three major food classes (carbohydrates, fats, and 


proteins), which is digested in the mouth, the stomach, and the small 
intestine? 


Solution: 


Answers may vary. 


Review Questions 


Exercise: 


Problem: 
In which part of the alimentary canal does most digestion occur? 


a. stomach 
b. proximal small intestine 


c. distal small intestine 
d. ascending colon 


Solution: 


B 


Exercise: 


Problem: Which of these is most associated with villi? 


a. haustra 

b. lacteals 

c. bacterial flora 

d. intestinal glands 


Solution: 
B 


Exercise: 


Problem: What is the role of the small intestine’s MALT? 


a. secreting mucus 

b. buffering acidic chyme 

c. activating pepsin 

d. preventing bacteria from entering the bloodstream 


Solution: 


D 


Exercise: 


Problem: Which part of the large intestine attaches to the appendix? 


a. cecum 
b. ascending colon 
c. transverse colon 
d. descending colon 


Solution: 


A 


Critical Thinking Questions 


Exercise: 
Problem: 


Explain how nutrients absorbed in the small intestine pass into the 
general circulation. 


Solution: 


Nutrients from the breakdown of carbohydrates and proteins are 
absorbed through a capillary bed in the villi of the small intestine. 
Lipid breakdown products are absorbed into a lacteal in the villi, and 
transported via the lymphatic system to the bloodstream. 


Exercise: 
Problem: 


Why is it important that chyme from the stomach is delivered to the 
small intestine slowly and in small amounts? 


Solution: 


If large quantities of chyme were forced into the small intestine, it 
would result in osmotic water loss from the blood into the intestinal 
lumen that could cause potentially life-threatening low blood volume 
and erosion of the duodenum. 


Exercise: 


Problem: 


Describe three of the differences between the walls of the large and 
small intestines. 


Solution: 


The mucosa of the small intestine includes circular folds, villi, and 
microvilli. The wall of the large intestine has a thick mucosal layer, 
and deeper and more abundant mucus-secreting glands that facilitate 
the smooth passage of feces. There are three features that are unique to 
the large intestine: teniae coli, haustra, and epiploic appendages. 


References 


American Cancer Society (US). Cancer facts and figures: colorectal cancer: 
2011-2013 [Internet]. c2013 [cited 2013 Apr 3]. Available from: 
http://www.cancer.org/Research/CancerFactsFigures/ColorectalCancerFacts 
Figures/colorectal-cancer-facts-figures-2011-2013-page. 


The Nutrition Source. Fiber and colon cancer: following the scientific trail 
[Internet]. Boston (MA): Harvard School of Public Health; c2012 [cited 
2013 Apr 3]. Available from: 
http://www.hsph.harvard.edu/nutritionsource/nutrition-news/fiber-and- 
colon-cancer/index. html. 


Centers for Disease Control and Prevention (US). Morbidity and mortality 
weekly report: notifiable diseases and mortality tables [Internet]. Atlanta 
(GA); [cited 2013 Apr 3]. Available from: 
http://www.cdc.gov/mmwr/preview/mmwrhtml/mm6101md.htm? 
s_cid=mm6101imd_w. 


Glossary 


anal canal 


final segment of the large intestine 


anal column 
long fold of mucosa in the anal canal 


anal sinus 
recess between anal columns 


appendix 
(vermiform appendix) coiled tube attached to the cecum 


ascending colon 
first region of the colon 


bacterial flora 
bacteria in the large intestine 


brush border 
fuzzy appearance of the small intestinal mucosa created by microvilli 


cecum 
pouch forming the beginning of the large intestine 


circular fold 
(also, plica circulare) deep fold in the mucosa and submucosa of the 
small intestine 


colon 
part of the large intestine between the cecum and the rectum 


descending colon 
part of the colon between the transverse colon and the sigmoid colon 


duodenal gland 
(also, Brunner’s gland) mucous-secreting gland in the duodenal 
submucosa 


duodenum 


first part of the small intestine, which starts at the pyloric sphincter and 
ends at the jejunum 


epiploic appendage 
small sac of fat-filled visceral peritoneum attached to teniae coli 


external anal sphincter 
voluntary skeletal muscle sphincter in the anal canal 


feces 
semisolid waste product of digestion 


flatus 
gas in the intestine 


gastrocolic reflex 
propulsive movement in the colon activated by the presence of food in 
the stomach 


gastroileal reflex 
long reflex that increases the strength of segmentation in the ileum 


haustrum 
small pouch in the colon created by tonic contractions of teniae coli 


haustral contraction 
slow segmentation in the large intestine 


hepatopancreatic ampulla 
(also, ampulla of Vater) bulb-like point in the wall of the duodenum 
where the bile duct and main pancreatic duct unite 


hepatopancreatic sphincter 
(also, sphincter of Oddi) sphincter regulating the flow of bile and 
pancreatic juice into the duodenum 


ileocecal sphincter 
sphincter located where the small intestine joins with the large 
intestine 


ileum 
end of the small intestine between the jejunum and the large intestine 


internal anal sphincter 
involuntary smooth muscle sphincter in the anal canal 


intestinal gland 
(also, crypt of Lieberktihn) gland in the small intestinal mucosa that 
secretes intestinal juice 


intestinal juice 
mixture of water and mucus that helps absorb nutrients from chyme 


jejunum 
middle part of the small intestine between the duodenum and the ileum 


lacteal 
lymphatic capillary in the villi 


large intestine 
terminal portion of the alimentary canal 


left colic flexure 
(also, splenic flexure) point where the transverse colon curves below 
the inferior end of the spleen 


main pancreatic duct 
(also, duct of Wirsung) duct through which pancreatic juice drains 
from the pancreas 


major duodenal papilla 
point at which the hepatopancreatic ampulla opens into the duodenum 


mass movement 
long, slow, peristaltic wave in the large intestine 


mesoappendix 
mesentery of the appendix 


microvillus 
small projection of the plasma membrane of the absorptive cells of the 
small intestinal mucosa 


migrating motility complex 
form of peristalsis in the small intestine 


motilin 
hormone that initiates migrating motility complexes 


pectinate line 
horizontal line that runs like a ring, perpendicular to the inferior 
margins of the anal sinuses 


rectal valve 
one of three transverse folds in the rectum where feces is separated 
from flatus 


rectum 
part of the large intestine between the sigmoid colon and anal canal 


right colic flexure 
(also, hepatic flexure) point, at the inferior surface of the liver, where 
the ascending colon turns abruptly to the left 


saccharolytic fermentation 
anaerobic decomposition of carbohydrates 


sigmoid colon 
end portion of the colon, which terminates at the rectum 


small intestine 
section of the alimentary canal where most digestion and absorption 
occurs 


tenia coli 
one of three smooth muscle bands that make up the longitudinal 
muscle layer of the muscularis in all of the large intestine except the 


terminal end 


transverse colon 
part of the colon between the ascending colon and the descending 
colon 


Valsalva’s maneuver 
voluntary contraction of the diaphragm and abdominal wall muscles 
and closing of the glottis, which increases intra-abdominal pressure 
and facilitates defecation 


villus 
projection of the mucosa of the small intestine 


Accessory Organs in Digestion: The Liver, Pancreas, and Gallbladder 
By the end of this section, you will be able to: 


e State the main digestive roles of the liver, pancreas, and gallbladder 
e Identify three main features of liver histology that are critical to its 
function 

Discuss the composition and function of bile 

Identify the major types of enzymes and buffers present in pancreatic 
juice 


Chemical digestion in the small intestine relies on the activities of three 
accessory digestive organs: the liver, pancreas, and gallbladder ({link]). The 
digestive role of the liver is to produce bile and export it to the duodenum. 
The gallbladder primarily stores, concentrates, and releases bile. The 
pancreas produces pancreatic juice, which contains digestive enzymes and 
bicarbonate ions, and delivers it to the duodenum. 

Accessory Organs 


Liver: 

Right lobe 
Quadrate lobe 
Left lobe 
Caudate lobe 


Gallbladder Spleen 


Right hepatic duct 


Pancreas 
Cystic duct 


Common hepatic duct Pancreatic duct 


Common bile duct 
Left hepatic duct 


The liver, pancreas, and gallbladder are 
considered accessory digestive organs, but 
their roles in the digestive system are vital. 


The Liver 


The liver is the largest gland in the body, weighing about three pounds in an 
adult. It is also one of the most important organs. In addition to being an 
accessory digestive organ, it plays a number of roles in metabolism and 
regulation. The liver lies inferior to the diaphragm in the right upper 
quadrant of the abdominal cavity and receives protection from the 
surrounding ribs. 


The liver is divided into two primary lobes: a large right lobe and a much 
smaller left lobe. In the right lobe, some anatomists also identify an inferior 
quadrate lobe and a posterior caudate lobe, which are defined by internal 
features. The liver is connected to the abdominal wall and diaphragm by 
five peritoneal folds referred to as ligaments. These are the falciform 
ligament, the coronary ligament, two lateral ligaments, and the ligamentum 
teres hepatis. The falciform ligament and ligamentum teres hepatis are 
actually remnants of the umbilical vein, and separate the right and left lobes 
anteriorly. The lesser omentum tethers the liver to the lesser curvature of the 
stomach. 


The porta hepatis (“gate to the liver”) is where the hepatic artery and 
hepatic portal vein enter the liver. These two vessels, along with the 
common hepatic duct, run behind the lateral border of the lesser omentum 
on the way to their destinations. As shown in [link], the hepatic artery 
delivers oxygenated blood from the heart to the liver. The hepatic portal 
vein delivers partially deoxygenated blood containing nutrients absorbed 
from the small intestine and actually supplies more oxygen to the liver than 
do the much smaller hepatic arteries. In addition to nutrients, drugs and 
toxins are also absorbed. After processing the bloodborne nutrients and 
toxins, the liver releases nutrients needed by other cells back into the blood, 
which drains into the central vein and then through the hepatic vein to the 
inferior vena cava. With this hepatic portal circulation, all blood from the 
alimentary canal passes through the liver. This largely explains why the 
liver is the most common site for the metastasis of cancers that originate in 
the alimentary canal. 

Microscopic Anatomy of the Liver 


Central vein 
Connective 
tissue 

Lobules 


Interlobular vein 
(to hepatic vein) 


Central vein Sinusoids 


Plates of Portal venule 
hepatocytes 


From portal vein 


The liver receives oxygenated blood from the 
hepatic artery and nutrient-rich deoxygenated 
blood from the hepatic portal vein. 


Histology 


The liver has three main components: hepatocytes, bile canaliculi, and 
hepatic sinusoids. A hepatocyte is the liver’s main cell type, accounting for 
around 80 percent of the liver's volume. These cells play a role in a wide 
variety of secretory, metabolic, and endocrine functions. Plates of 


hepatocytes called hepatic laminae radiate outward from the portal vein in 
each hepatic lobule. 


Between adjacent hepatocytes, grooves in the cell membranes provide room 
for each bile canaliculus (plural = canaliculi). These small ducts 
accumulate the bile produced by hepatocytes. From here, bile flows first 
into bile ductules and then into bile ducts. The bile ducts unite to form the 
larger right and left hepatic ducts, which themselves merge and exit the 
liver as the common hepatic duct. This duct then joins with the cystic duct 
from the gallbladder, forming the common bile duct through which bile 
flows into the small intestine. 


A hepatic sinusoid is an open, porous blood space formed by fenestrated 
capillaries from nutrient-rich hepatic portal veins and oxygen-rich hepatic 
arteries. Hepatocytes are tightly packed around the fenestrated endothelium 
of these spaces, giving them easy access to the blood. From their central 
position, hepatocytes process the nutrients, toxins, and waste materials 
carried by the blood. Materials such as bilirubin are processed and excreted 
into the bile canaliculi. Other materials including proteins, lipids, and 
carbohydrates are processed and secreted into the sinusoids or just stored in 
the cells until called upon. The hepatic sinusoids combine and send blood to 
a central vein. Blood then flows through a hepatic vein into the inferior 
vena cava. This means that blood and bile flow in opposite directions. The 
hepatic sinusoids also contain star-shaped reticuloendothelial cells 
(Kupffer cells), phagocytes that remove dead red and white blood cells, 
bacteria, and other foreign material that enter the sinusoids. The portal 
triad is a distinctive arrangement around the perimeter of hepatic lobules, 
consisting of three basic structures: a bile duct, a hepatic artery branch, and 
a hepatic portal vein branch. 


Bile 


Recall that lipids are hydrophobic, that is, they do not dissolve in water. 
Thus, before they can be digested in the watery environment of the small 
intestine, large lipid globules must be broken down into smaller lipid 


globules, a process called emulsification. Bile is a mixture secreted by the 
liver to accomplish the emulsification of lipids in the small intestine. 


Hepatocytes secrete about one liter of bile each day. A yellow-brown or 
yellow-green alkaline solution (pH 7.6 to 8.6), bile is a mixture of water, 
bile salts, bile pigments, phospholipids (such as lecithin), electrolytes, 
cholesterol, and triglycerides. The components most critical to 
emulsification are bile salts and phospholipids, which have a nonpolar 
(hydrophobic) region as well as a polar (hydrophilic) region. The 
hydrophobic region interacts with the large lipid molecules, whereas the 
hydrophilic region interacts with the watery chyme in the intestine. This 
results in the large lipid globules being pulled apart into many tiny lipid 
fragments of about 1 ym in diameter. This change dramatically increases 
the surface area available for lipid-digesting enzyme activity. This is the 
same way dish soap works on fats mixed with water. 


Bile salts act as emulsifying agents, so they are also important for the 
absorption of digested lipids. While most constituents of bile are eliminated 
in feces, bile salts are reclaimed by the enterohepatic circulation. Once 
bile salts reach the ileum, they are absorbed and returned to the liver in the 
hepatic portal blood. The hepatocytes then excrete the bile salts into newly 
formed bile. Thus, this precious resource is recycled. 


Bilirubin, the main bile pigment, is a waste product produced when the 
spleen removes old or damaged red blood cells from the circulation. These 
breakdown products, including proteins, iron, and toxic bilirubin, are 
transported to the liver via the splenic vein of the hepatic portal system. In 
the liver, proteins and iron are recycled, whereas bilirubin is excreted in the 
bile. It accounts for the green color of bile. Bilirubin is eventually 
transformed by intestinal bacteria into stercobilin, a brown pigment that 
gives your stool its characteristic color! In some disease states, bile does not 
enter the intestine, resulting in white (‘acholic’) stool with a high fat 
content, since virtually no fats are broken down or absorbed. 


Hepatocytes work non-stop, but bile production increases when fatty chyme 
enters the duodenum and stimulates the secretion of the gut hormone 
secretin. Between meals, bile is produced but conserved. The valve-like 


hepatopancreatic ampulla closes, allowing bile to divert to the gallbladder, 
where it is concentrated and stored until the next meal. 


Note: 


. . 
mss’ OPENStax COLLEGE 


Watch this video to see the structure of the liver and how this structure 
supports the functions of the liver, including the processing of nutrients, 
toxins, and wastes. At rest, about 1500 mL of blood per minute flow 
through the liver. What percentage of this blood flow comes from the 
hepatic portal system? 


The Pancreas 


The soft, oblong, glandular pancreas lies transversely in the 
retroperitoneum behind the stomach. Its head is nestled into the “c-shaped” 
curvature of the duodenum with the body extending to the left about 15.2 
cm (6 in) and ending as a tapering tail in the hilum of the spleen. It is a 
curious mix of exocrine (secreting digestive enzymes) and endocrine 
(releasing hormones into the blood) functions ([link]). 

Exocrine and Endocrine Pancreas 


Common bile duct Pancreatic duct 


Tail of 
pancreas 


Lobules 


Acinar cells secrete 
Head of pancreas digestive enzymes. 


Pancreatic islet) _ ® Sie as 
cells secrete We = 
hormones. ‘@ 


Pancreatic duct 


Exocrine cells secrete pancreatic juice. 


The pancreas has a head, a body, and a 
tail. It delivers pancreatic juice to the 
duodenum through the pancreatic 
duct. 


The exocrine part of the pancreas arises as little grape-like cell clusters, 
each called an acinus (plural = acini), located at the terminal ends of 
pancreatic ducts. These acinar cells secrete enzyme-rich pancreatic juice 
into tiny merging ducts that form two dominant ducts. The larger duct fuses 
with the common bile duct (carrying bile from the liver and gallbladder) 
just before entering the duodenum via a common opening (the 
hepatopancreatic ampulla). The smooth muscle sphincter of the 
hepatopancreatic ampulla controls the release of pancreatic juice and bile 
into the small intestine. The second and smaller pancreatic duct, the 


accessory duct (duct of Santorini), runs from the pancreas directly into the 
duodenum, approximately 1 inch above the hepatopancreatic ampulla. 
When present, it is a persistent remnant of pancreatic development. 


Scattered through the sea of exocrine acini are small islands of endocrine 
cells, the islets of Langerhans. These vital cells produce the hormones 
pancreatic polypeptide, insulin, glucagon, and somatostatin. 


Pancreatic Juice 


The pancreas produces over a liter of pancreatic juice each day. Unlike bile, 
it is clear and composed mostly of water along with some salts, sodium 
bicarbonate, and several digestive enzymes. Sodium bicarbonate is 
responsible for the slight alkalinity of pancreatic juice (pH 7.1 to 8.2), 
which serves to buffer the acidic gastric juice in chyme, inactivate pepsin 
from the stomach, and create an optimal environment for the activity of pH- 
sensitive digestive enzymes in the small intestine. Pancreatic enzymes are 
active in the digestion of sugars, proteins, and fats. 


The pancreas produces protein-digesting enzymes in their inactive forms. 
These enzymes are activated in the duodenum. If produced in an active 
form, they would digest the pancreas (which is exactly what occurs in the 
disease, pancreatitis). The intestinal brush border enzyme enteropeptidase 
stimulates the activation of trypsin from trypsinogen of the pancreas, which 
in turn changes the pancreatic enzymes procarboxypeptidase and 
chymotrypsinogen into their active forms, carboxypeptidase and 
chymotrypsin. 


The enzymes that digest starch (amylase), fat (lipase), and nucleic acids 
(nuclease) are secreted in their active forms, since they do not attack the 
pancreas as do the protein-digesting enzymes. 


Pancreatic Secretion 


Regulation of pancreatic secretion is the job of hormones and the 
parasympathetic nervous system. The entry of acidic chyme into the 
duodenum stimulates the release of secretin, which in turn causes the duct 
cells to release bicarbonate-rich pancreatic juice. The presence of proteins 
and fats in the duodenum stimulates the secretion of CCK, which then 
stimulates the acini to secrete enzyme-rich pancreatic juice and enhances 
the activity of secretin. Parasympathetic regulation occurs mainly during 
the cephalic and gastric phases of gastric secretion, when vagal stimulation 
prompts the secretion of pancreatic juice. 


Usually, the pancreas secretes just enough bicarbonate to counterbalance the 
amount of HCl produced in the stomach. Hydrogen ions enter the blood 
when bicarbonate is secreted by the pancreas. Thus, the acidic blood 
draining from the pancreas neutralizes the alkaline blood draining from the 
stomach, maintaining the pH of the venous blood that flows to the liver. 


The Gallbladder 


The gallbladder is 8—10 cm (~3-4 in) long and is nested in a shallow area 
on the posterior aspect of the right lobe of the liver. This muscular sac 
stores, concentrates, and, when stimulated, propels the bile into the 
duodenum via the common bile duct. It is divided into three regions. The 
fundus is the widest portion and tapers medially into the body, which in turn 
narrows to become the neck. The neck angles slightly superiorly as it 
approaches the hepatic duct. The cystic duct is 1—2 cm (less than 1 in) long 
and turns inferiorly as it bridges the neck and hepatic duct. 


The simple columnar epithelium of the gallbladder mucosa is organized in 
rugae, similar to those of the stomach. There is no submucosa in the 
gallbladder wall. The wall’s middle, muscular coat is made of smooth 
muscle fibers. When these fibers contract, the gallbladder’s contents are 
ejected through the cystic duct and into the bile duct ([link]). Visceral 
peritoneum reflected from the liver capsule holds the gallbladder against the 
liver and forms the outer coat of the gallbladder. The gallbladder's mucosa 
absorbs water and ions from bile, concentrating it by up to 10-fold. 
Gallbladder 


Left hepatic duct 
Right hepatic duct 
Cystic duct 


Gallbladder: 
Body 
Fundus 
Neck 


Common 
hepatic duct 


F Common 
Liver bile duct 


The gallbladder stores and concentrates 
bile, and releases it into the two-way 
cystic duct when it is needed by the small 
intestine. 


Chapter Review 


Chemical digestion in the small intestine cannot occur without the help of 
the liver and pancreas. The liver produces bile and delivers it to the 
common hepatic duct. Bile contains bile salts and phospholipids, which 
emulsify large lipid globules into tiny lipid droplets, a necessary step in 
lipid digestion and absorption. The gallbladder stores and concentrates bile, 
releasing it when it is needed by the small intestine. 


The pancreas produces the enzyme- and bicarbonate-rich pancreatic juice 
and delivers it to the small intestine through ducts. Pancreatic juice buffers 
the acidic gastric juice in chyme, inactivates pepsin from the stomach, and 
enables the optimal functioning of digestive enzymes in the small intestine. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to see the structure of the liver and how this structure 
supports the functions of the liver, including the processing of 
nutrients, toxins, and wastes. At rest, about 1500 mL of blood per 
minute flow through the liver. What percentage of this blood flow 
comes from the hepatic portal system? 


Solution: 


Answers may vary. 


Review Questions 


Exercise: 


Problem: Which of these statements about bile is true? 


a. About 500 mL is secreted daily. 

b. Its main function is the denaturation of proteins. 
c. It is synthesized in the gallbladder. 

d. Bile salts are recycled. 


Solution: 


D 


Exercise: 


Problem: Pancreatic juice 


a. deactivates bile. 

b. is secreted by pancreatic islet cells. 
c. buffers chyme. 

d. is released into the cystic duct. 


Solution: 


C 


Critical Thinking Questions 


Exercise: 
Problem: 


Why does the pancreas secrete some enzymes in their inactive forms, 
and where are these enzymes activated? 


Solution: 


The pancreas secretes protein-digesting enzymes in their inactive 
forms. If secreted in their active forms, they would self-digest the 
pancreas. These enzymes are activated in the duodenum. 


Exercise: 


Problem: 


Describe the location of hepatocytes in the liver and how this 
arrangement enhances their function. 


Solution: 


The hepatocytes are the main cell type of the liver. They process, store, 
and release nutrients into the blood. Radiating out from the central 
vein, they are tightly packed around the hepatic sinusoids, allowing the 
hepatocytes easy access to the blood flowing through the sinusoids. 


Glossary 


accessory duct 


(also, duct of Santorini) duct that runs from the pancreas into the 
duodenum 


acinus 
cluster of glandular epithelial cells in the pancreas that secretes 
pancreatic juice in the pancreas 


bile 
alkaline solution produced by the liver and important for the 
emulsification of lipids 


bile canaliculus 
small duct between hepatocytes that collects bile 


bilirubin 
main bile pigment, which is responsible for the brown color of feces 


central vein 
vein that receives blood from hepatic sinusoids 


common bile duct 
structure formed by the union of the common hepatic duct and the 
gallbladder’s cystic duct 


common hepatic duct 
duct formed by the merger of the two hepatic ducts 


cystic duct 
duct through which bile drains and enters the gallbladder 


enterohepatic circulation 
recycling mechanism that conserves bile salts 


enteropeptidase 
intestinal brush-border enzyme that activates trypsinogen to trypsin 


gallbladder 
accessory digestive organ that stores and concentrates bile 


hepatic artery 
artery that supplies oxygenated blood to the liver 


hepatic lobule 
hexagonal-shaped structure composed of hepatocytes that radiate 
outward from a central vein 


hepatic portal vein 
vein that supplies deoxygenated nutrient-rich blood to the liver 


hepatic sinusoid 
blood capillaries between rows of hepatocytes that receive blood from 
the hepatic portal vein and the branches of the hepatic artery 


hepatic vein 
vein that drains into the inferior vena cava 


hepatocytes 
major functional cells of the liver 


liver 
largest gland in the body whose main digestive function is the 
production of bile 


pancreas 
accessory digestive organ that secretes pancreatic juice 


pancreatic juice 
secretion of the pancreas containing digestive enzymes and 
bicarbonate 


porta hepatis 
“gateway to the liver” where the hepatic artery and hepatic portal vein 


enter the liver 


portal triad 
bile duct, hepatic artery branch, and hepatic portal vein branch 


reticuloendothelial cell 


(also, Kupffer cell) phagocyte in hepatic sinusoids that filters out 
material from venous blood from the alimentary canal 


Anatomy of the Lymphatic and Immune Systems 
By the end of this section, you will be able to: 


e Describe the structure and function of the lymphatic tissue (lymph 
fluid, vessels, ducts, and organs) 

e Describe the structure and function of the primary and secondary 
lymphatic organs 

e Discuss the cells of the immune system, how they function, and their 
relationship with the lymphatic system 


The immune system is the complex collection of cells and organs that 
destroys or neutralizes pathogens that would otherwise cause disease or 
death. The lymphatic system, for most people, is associated with the 
immune system to such a degree that the two systems are virtually 
indistinguishable. The lymphatic system is the system of vessels, cells, and 
organs that carries excess fluids to the bloodstream and filters pathogens 
from the blood. The swelling of lymph nodes during an infection and the 
transport of lymphocytes via the lymphatic vessels are but two examples of 
the many connections between these critical organ systems. 


Functions of the Lymphatic System 


A major function of the lymphatic system is to drain body fluids and return 
them to the bloodstream. Blood pressure causes leakage of fluid from the 
capillaries, resulting in the accumulation of fluid in the interstitial space— 
that is, spaces between individual cells in the tissues. In humans, 20 liters of 
plasma is released into the interstitial space of the tissues each day due to 
capillary filtration. Once this filtrate is out of the bloodstream and in the 
tissue spaces, it is referred to as interstitial fluid. Of this, 17 liters is 
reabsorbed directly by the blood vessels. But what happens to the remaining 
three liters? This is where the lymphatic system comes into play. It drains 
the excess fluid and empties it back into the bloodstream via a series of 
vessels, trunks, and ducts. Lymph is the term used to describe interstitial 
fluid once it has entered the lymphatic system. When the lymphatic system 
is damaged in some way, such as by being blocked by cancer cells or 
destroyed by injury, protein-rich interstitial fluid accumulates (sometimes 
“backs up” from the lymph vessels) in the tissue spaces. This inappropriate 


accumulation of fluid referred to as lymphedema may lead to serious 
medical consequences. 


As the vertebrate immune system evolved, the network of lymphatic vessels 
became convenient avenues for transporting the cells of the immune 
system. Additionally, the transport of dietary lipids and fat-soluble vitamins 
absorbed in the gut uses this system. 


Cells of the immune system not only use lymphatic vessels to make their 
way from interstitial spaces back into the circulation, but they also use 
lymph nodes as major staging areas for the development of critical immune 
responses. A lymph node is one of the small, bean-shaped organs located 
throughout the lymphatic system. 


Note: 


openstax COLLEGE” 


Visit this website for an overview of the lymphatic system. What are the 
three main components of the lymphatic system? 


Structure of the Lymphatic System 


The lymphatic vessels begin as open-ended capillaries, which feed into 
larger and larger lymphatic vessels, and eventually empty into the 
bloodstream by a series of ducts. Along the way, the lymph travels through 
the lymph nodes, which are commonly found near the groin, armpits, neck, 
chest, and abdomen. Humans have about 500—600 lymph nodes throughout 
the body ([link]). 

Anatomy of the Lymphatic System 


Adenoid 
Tonsil 


J 
Ah Lymph nodes 


_ j 4\ Thymus 
| Lymphatic 


| vessel 


; 7 Thymus 
Right lymphatic duct, 
entering vein 


Tissue cell 


Spleen 


Interstitial 
fluid 


capillary ot 


Lymphatic 
capillary 


Masses of lymphocytes 
and macrophages 


Bone marrow 


Lymph node 


Lymphatic vessels in the arms and legs convey lymph 
to the larger lymphatic vessels in the torso. 


A major distinction between the lymphatic and cardiovascular systems in 
humans is that lymph is not actively pumped by the heart, but is forced 
through the vessels by the movements of the body, the contraction of 
skeletal muscles during body movements, and breathing. One-way valves 
(semi-lunar valves) in lymphatic vessels keep the lymph moving toward the 
heart. Lymph flows from the lymphatic capillaries, through lymphatic 
vessels, and then is dumped into the circulatory system via the lymphatic 
ducts located at the junction of the jugular and subclavian veins in the neck. 


Lymphatic Capillaries 


Lymphatic capillaries, also called the terminal lymphatics, are vessels 
where interstitial fluid enters the lymphatic system to become lymph fluid. 
Located in almost every tissue in the body, these vessels are interlaced 
among the arterioles and venules of the circulatory system in the soft 
connective tissues of the body ({link]). Exceptions are the central nervous 
system, bone marrow, bones, teeth, and the cornea of the eye, which do not 
contain lymph vessels. 

Lymphatic Capillaries 


Lymph capillaries in the tissue spaces 


Lymph capillary. 
Gy x Collagen fiber 


Arteriole 
Interstitial fluid 


Lymph (interstitial fluid) 


x Lymphatic 


vessel 


Tissue fluid 
Endothelial 
“flaps” 
Lymph vessel 

endothelial cells 


prevention 
valve 


Lymphatic capillaries are interlaced with the arterioles 
and venules of the cardiovascular system. Collagen 
fibers anchor a lymphatic capillary in the tissue (inset). 
Interstitial fluid slips through spaces between the 
overlapping endothelial cells that compose the lymphatic 
capillary. 


Lymphatic capillaries are formed by a one cell-thick layer of endothelial 
cells and represent the open end of the system, allowing interstitial fluid to 
flow into them via overlapping cells (see [link]). When interstitial pressure 
is low, the endothelial flaps close to prevent “backflow.” As interstitial 
pressure increases, the spaces between the cells open up, allowing the fluid 


to enter. Entry of fluid into lymphatic capillaries is also enabled by the 
collagen filaments that anchor the capillaries to surrounding structures. As 
interstitial pressure increases, the filaments pull on the endothelial cell 
flaps, opening up them even further to allow easy entry of fluid. 


In the small intestine, lymphatic capillaries called lacteals are critical for the 
transport of dietary lipids and lipid-soluble vitamins to the bloodstream. In 
the small intestine, dietary triglycerides combine with other lipids and 
proteins, and enter the lacteals to form a milky fluid called chyle. The chyle 
then travels through the lymphatic system, eventually entering the 
bloodstream. 


Larger Lymphatic Vessels, Trunks, and Ducts 


The lymphatic capillaries empty into larger lymphatic vessels, which are 
similar to veins in terms of their three-tunic structure and the presence of 
valves. These one-way valves are located fairly close to one another, and 
each one causes a bulge in the lymphatic vessel, giving the vessels a beaded 
appearance (see [link]). 


The superficial and deep lymphatics eventually merge to form larger 
lymphatic vessels known as lymphatic trunks. On the right side of the 
body, the right sides of the head, thorax, and right upper limb drain lymph 
fluid into the right subclavian vein via the right lymphatic duct ([link]). On 
the left side of the body, the remaining portions of the body drain into the 
larger thoracic duct, which drains into the left subclavian vein. The thoracic 
duct itself begins just beneath the diaphragm in the cisterna chyli, a sac- 
like chamber that receives lymph from the lower abdomen, pelvis, and 
lower limbs by way of the left and right lumbar trunks and the intestinal 
trunk. 

Major Trunks and Ducts of the Lymphatic System 


Right lymphatic Right internal Left internal Thoracic duct 
duct jugular vein jugular vein drains into 


subclavian vein 


Left subclavian vein 


Right subclavian vein 


Thoracic duct 


Cisterna chyli of 


Drained by right thoracic duct 


lymphatic duct 


Drained by 
thoracic duct 


The thoracic duct drains a much larger portion of the 
body than does the right lymphatic duct. 


The overall drainage system of the body is asymmetrical (see [link]). The 
right lymphatic duct receives lymph from only the upper right side of the 
body. The lymph from the rest of the body enters the bloodstream through 
the thoracic duct via all the remaining lymphatic trunks. In general, 
lymphatic vessels of the subcutaneous tissues of the skin, that is, the 
superficial lymphatics, follow the same routes as veins, whereas the deep 
lymphatic vessels of the viscera generally follow the paths of arteries. 


The Organization of Immune Function 


The immune system is a collection of barriers, cells, and soluble proteins 
that interact and communicate with each other in extraordinarily complex 
ways. The modern model of immune function is organized into three phases 


based on the timing of their effects. The three temporal phases consist of the 
following: 


¢ Barrier defenses such as the skin and mucous membranes, which act 
instantaneously to prevent pathogenic invasion into the body tissues 

e The rapid but nonspecific innate immune response, which consists of 
a variety of specialized cells and soluble factors 

e The slower but more specific and effective adaptive immune 
response, which involves many cell types and soluble factors, but is 
primarily controlled by white blood cells (leukocytes) known as 
lymphocytes, which help control immune responses 


The cells of the blood, including all those involved in the immune response, 
arise in the bone marrow via various differentiation pathways from 
hematopoietic stem cells ({link]). In contrast with embryonic stem cells, 
hematopoietic stem cells are present throughout adulthood and allow for the 
continuous differentiation of blood cells to replace those lost to age or 
function. These cells can be divided into three classes based on function: 


e Phagocytic cells, which ingest pathogens to destroy them 

e Lymphocytes, which specifically coordinate the activities of adaptive 
immunity 

¢ Cells containing cytoplasmic granules, which help mediate immune 
responses against parasites and intracellular pathogens such as viruses 


Hematopoietic System of the Bone Marrow 


After division some cells 
remain stem cells. 


Multipotent hematopoietic 
stem cell (hemocytoblast) 


a 
e 


, The remaining cell goes down one of two paths 
@ depending on the chemical signals received. 


e 
Myeloid stem cell Lymphoid stem cell 


Megakaryoblast Proerythroblast Myeloblast Monoblast Lymphoblast 


@ 


Reticulocyte 


=i oo 
yy rc ae Le Small lymphocyte 
ork A 2 : —_— arge granular 

Mm @ ovr YY 

| — “ 
Megakaryocyte Erythrocyte Basophil Neutrophil Eosinophil Monocyte sy) @& 

T lymphocyte —_B lymphocyte 

b) 
p) 


Ad) 
BY) 
7 


7X SS 
Platelets 7) => o 


° Plasma cell 
Macrophage 


All the cells of the immune response as well as of the blood arise by 
differentiation from hematopoietic stem cells. Platelets are cell 
fragments involved in the clotting of blood. 


Lymphocytes: B Cells, T Cells, Plasma Cells, and Natural 
Killer Cells 


As stated above, lymphocytes are the primary cells of adaptive immune 
responses ({link]). The two basic types of lymphocytes, B cells and T cells, 
are identical morphologically with a large central nucleus surrounded by a 
thin layer of cytoplasm. They are distinguished from each other by their 
surface protein markers as well as by the molecules they secrete. While B 


cells mature in red bone marrow and T cells mature in the thymus, they 
both initially develop from bone marrow. T cells migrate from bone marrow 
to the thymus gland where they further mature. B cells and T cells are found 
in many parts of the body, circulating in the bloodstream and lymph, and 
residing in secondary lymphoid organs, including the spleen and lymph 
nodes, which will be described later in this section. The human body 
contains approximately 10'* lymphocytes. 


B Cells 


B cells are immune cells that function primarily by producing antibodies. 
An antibody is any of the group of proteins that binds specifically to 
pathogen-associated molecules known as antigens. An antigen is a 
chemical structure on the surface of a pathogen that binds to T or B 
lymphocyte antigen receptors. Once activated by binding to antigen, B cells 
differentiate into cells that secrete a soluble form of their surface antibodies. 
These activated B cells are known as plasma cells. 


T Cells 


The T cell, on the other hand, does not secrete antibody but performs a 
variety of functions in the adaptive immune response. Different T cell types 
have the ability to either secrete soluble factors that communicate with 
other cells of the adaptive immune response or destroy cells infected with 
intracellular pathogens. The roles of T and B lymphocytes in the adaptive 
immune response will be discussed further in this chapter. 


Plasma Cells 


Another type of lymphocyte of importance is the plasma cell. A plasma cell 
is a B cell that has differentiated in response to antigen binding, and has 
thereby gained the ability to secrete soluble antibodies. These cells differ in 
morphology from standard B and T cells in that they contain a large amount 


of cytoplasm packed with the protein-synthesizing machinery known as 
rough endoplasmic reticulum. 


Natural Killer Cells 


A fourth important lymphocyte is the natural killer cell, a participant in the 
innate immune response. A natural killer cell (NK) is a circulating blood 
cell that contains cytotoxic (cell-killing) granules in its extensive 
cytoplasm. It shares this mechanism with the cytotoxic T cells of the 
adaptive immune response. NK cells are among the body’s first lines of 
defense against viruses and certain types of cancer. 


Lymphocytes 

Type of lymphocyte Primary function 

B lymphocyte Generates diverse antibodies 
T lymphocyte Secretes chemical messengers 
Plasma cell Secretes antibodies 

NK cell Destroys virally infected cells 


Note: 


Visit this website to learn about the many different cell types in the 
immune system and their very specialized jobs. What is the role of the 
dendritic cell in an HIV infection? 


Primary Lymphoid Organs and Lymphocyte Development 


Understanding the differentiation and development of B and T cells is 
critical to the understanding of the adaptive immune response. It is through 
this process that the body (ideally) learns to destroy only pathogens and 
leaves the body’s own cells relatively intact. The primary lymphoid 
organs are the bone marrow and thymus gland. The lymphoid organs are 
where lymphocytes mature, proliferate, and are selected, which enables 
them to attack pathogens without harming the cells of the body. 


Bone Marrow 


In the embryo, blood cells are made in the yolk sac. As development 
proceeds, this function is taken over by the spleen, lymph nodes, and liver. 
Later, the bone marrow takes over most hematopoietic functions, although 
the final stages of the differentiation of some cells may take place in other 
organs. The red bone marrow is a loose collection of cells where 
hematopoiesis occurs, and the yellow bone marrow is a site of energy 
storage, which consists largely of fat cells ({link]). The B cell undergoes 
nearly all of its development in the red bone marrow, whereas the immature 
T cell, called a thymocyte, leaves the bone marrow and matures largely in 
the thymus gland. 

Bone Marrow 


Red bone marrow fills the head 


of the femur, and a spot of 
yellow bone marrow is visible 
in the center. The white 
reference bar is 1 cm. 


Thymus 


The thymus gland is a bilobed organ found in the space between the 
sternum and the aorta of the heart ({link]). Connective tissue holds the lobes 
closely together but also separates them and forms a capsule. 

Location, Structure, and Histology of the Thymus 


Cortex Trabeculae 


Fibrous 
capsule 


Right lymphatic duct, 
entering vein 


Lymph nodes 


Cortical epithelial cell Thymocytes _Trabecula 


Heart 
FONG oy Hai eee —}— Fibrous 
Oma \ Wows ke capsule 
3 5, e: . Cortex 
t <9 
Spleen A a Yo a Medulla 
Dendritic cell », ry. t 
Macrophage Blood vessel Medullary 
epithelial cell 


The thymus lies above the heart. The trabeculae and 
lobules, including the darkly staining cortex and the 
lighter staining medulla of each lobule, are clearly 
visible in the light micrograph of the thymus of a 
newborn. LM x 100. (Micrograph provided by the 
Regents of the University of Michigan Medical School 
© 2012) 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. 


The connective tissue capsule further divides the thymus into lobules via 
extensions called trabeculae. The outer region of the organ is known as the 
cortex and contains large numbers of thymocytes with some epithelial cells, 
macrophages, and dendritic cells (two types of phagocytic cells that are 
derived from monocytes). The cortex is densely packed so it stains more 
intensely than the rest of the thymus (see [link]). The medulla, where 
thymocytes migrate before leaving the thymus, contains a less dense 
collection of thymocytes, epithelial cells, and dendritic cells. 


Note: 

Aging and the... 

Immune System 

By the year 2050, 25 percent of the population of the United States will be 
60 years of age or older. The CDC estimates that 80 percent of those 60 
years and older have one or more chronic disease associated with 
deficiencies of the immune systems. This loss of immune function with age 
is called immunosenescence. To treat this growing population, medical 
professionals must better understand the aging process. One major cause of 
age-related immune deficiencies is thymic involution, the shrinking of the 
thymus gland that begins at birth, at a rate of about three percent tissue loss 
per year, and continues until 35-45 years of age, when the rate declines to 
about one percent loss per year for the rest of one’s life. At that pace, the 
total loss of thymic epithelial tissue and thymocytes would occur at about 
120 years of age. Thus, this age is a theoretical limit to a healthy human 
lifespan. 

Thymic involution has been observed in all vertebrate species that have a 
thymus gland. Animal studies have shown that transplanted thymic grafts 
between inbred strains of mice involuted according to the age of the donor 
and not of the recipient, implying the process is genetically programmed. 
There is evidence that the thymic microenvironment, so vital to the 


development of naive T cells, loses thymic epithelial cells according to the 
decreasing expression of the FOXNI1 gene with age. 

It is also known that thymic involution can be altered by hormone levels. 
Sex hormones such as estrogen and testosterone enhance involution, and 
the hormonal changes in pregnant women cause a temporary thymic 
involution that reverses itself, when the size of the thymus and its hormone 
levels return to normal, usually after lactation ceases. What does all this 
tell us? Can we reverse immunosenescence, or at least slow it down? The 
potential is there for using thymic transplants from younger donors to keep 
thymic output of naive T cells high. Gene therapies that target gene 
expression are also seen as future possibilities. The more we learn through 
immunosenescence research, the more opportunities there will be to 
develop therapies, even though these therapies will likely take decades to 
develop. The ultimate goal is for everyone to live and be healthy longer, 
but there may be limits to immortality imposed by our genes and 
hormones. 


Secondary Lymphoid Organs and their Roles in Active 
Immune Responses 


Lymphocytes develop and mature in the primary lymphoid organs, but they 
mount immune responses from the secondary lymphoid organs. A naive 
lymphocyte is one that has left the primary organ and entered a secondary 
lymphoid organ. Naive lymphocytes are fully functional immunologically, 
but have yet to encounter an antigen to respond to. In addition to circulating 
in the blood and lymph, lymphocytes concentrate in secondary lymphoid 
organs, which include the lymph nodes, spleen, and lymphoid nodules. All 
of these tissues have many features in common, including the following: 


e The presence of lymphoid follicles, the sites of the formation of 
lymphocytes, with specific B cell-rich and T cell-rich areas 

e An internal structure of reticular fibers with associated fixed 
macrophages 

¢ Germinal centers, which are the sites of rapidly dividing and 
differentiating B lymphocytes 


e Specialized post-capillary vessels known as high endothelial venules; 
the cells lining these venules are thicker and more columnar than 
normal endothelial cells, which allow cells from the blood to directly 
enter these tissues 


Lymph Nodes 


Lymph nodes function to remove debris and pathogens from the lymph, and 
are thus sometimes referred to as the “filters of the lymph” ({link]). Any 
bacteria that infect the interstitial fluid are taken up by the lymphatic 
capillaries and transported to a regional lymph node. Dendritic cells and 
macrophages within this organ internalize and kill many of the pathogens 
that pass through, thereby removing them from the body. The lymph node is 
also the site of adaptive immune responses mediated by T cells, B cells, and 
accessory Cells of the adaptive immune system. Like the thymus, the bean- 
shaped lymph nodes are surrounded by a tough capsule of connective tissue 
and are separated into compartments by trabeculae, the extensions of the 
capsule. In addition to the structure provided by the capsule and trabeculae, 
the structural support of the lymph node is provided by a series of reticular 
fibers laid down by fibroblasts. 

Structure and Histology of a Lymph Node 

Efferent lymphatic 


Connective tissue Cortex vessels 
capsule 


Ofc Z 
. J 
ie : Rive. 
fad | 2 Prats 
fe 2, 
~~ Roh . ‘ 
ey 3 : 
7 ‘ 3 ~ 
i 
- 


Connective 
tissue capsule 


Subcapsular 


Subcapsular 
sinus Afferent lymphatic vessels 


Lymph nodes are masses of lymphatic tissue located 
along the larger lymph vessels. The micrograph of the 
lymph nodes shows a germinal center, which consists of 
rapidly dividing B cells surrounded by a layer of T cells 
and other accessory cells. LM x 128. (Micrograph 


provided by the Regents of the University of Michigan 
Medical School © 2012) 


Note: 

[=] [a 

ro 

= pense COLLEGE 
. rg 

Ott lt 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. 


The major routes into the lymph node are via afferent lymphatic vessels 
(see [link]). Cells and lymph fluid that leave the lymph node may do so by 
another set of vessels known as the efferent lymphatic vessels. Lymph 
enters the lymph node via the subcapsular sinus, which is occupied by 
dendritic cells, macrophages, and reticular fibers. Within the cortex of the 
lymph node are lymphoid follicles, which consist of germinal centers of 
rapidly dividing B cells surrounded by a layer of T cells and other accessory 
cells. As the lymph continues to flow through the node, it enters the 
medulla, which consists of medullary cords of B cells and plasma cells, and 
the medullary sinuses where the lymph collects before leaving the node via 
the efferent lymphatic vessels. 


Spleen 


In addition to the lymph nodes, the spleen is a major secondary lymphoid 
organ ({link]). It is about 12 cm (5 in) long and is attached to the lateral 


border of the stomach via the gastrosplenic ligament. The spleen is a fragile 
organ without a strong capsule, and is dark red due to its extensive 
vascularization. The spleen is sometimes called the “filter of the blood” 
because of its extensive vascularization and the presence of macrophages 
and dendritic cells that remove microbes and other materials from the 
blood, including dying red blood cells. The spleen also functions as the 
location of immune responses to blood-borne pathogens. 

Spleen 


(a) Cross section of the spleen Hilum 


Trabecula 


Splenic vein 


Diaphragm White pulp Arteriole Venule 


Trabecula 


Marginal zone 


Central artery or 
arteriole 


= Germinal center 
Arterial capillaries =< = 2S 


2 
at 
Bp 4 


Venous sinus 


(a) The spleen is attached to the stomach. (b) A 
micrograph of spleen tissue shows the germinal center. 
The marginal zone is the region between the red pulp and 
white pulp, which sequesters particulate antigens from 
the circulation and presents these antigens to 
lymphocytes in the white pulp. EM x 660. (Micrograph 


provided by the Regents of the University of Michigan 
Medical School © 2012) 


The spleen is also divided by trabeculae of connective tissue, and within 
each splenic nodule is an area of red pulp, consisting of mostly red blood 
cells, and white pulp, which resembles the lymphoid follicles of the lymph 
nodes. Upon entering the spleen, the splenic artery splits into several 
arterioles (surrounded by white pulp) and eventually into sinusoids. Blood 
from the capillaries subsequently collects in the venous sinuses and leaves 
via the splenic vein. The red pulp consists of reticular fibers with fixed 
macrophages attached, free macrophages, and all of the other cells typical 
of the blood, including some lymphocytes. The white pulp surrounds a 
central arteriole and consists of germinal centers of dividing B cells 
surrounded by T cells and accessory cells, including macrophages and 
dendritic cells. Thus, the red pulp primarily functions as a filtration system 
of the blood, using cells of the relatively nonspecific immune response, and 
white pulp is where adaptive T and B cell responses are mounted. 


Lymphoid Nodules 


The other lymphoid tissues, the lymphoid nodules, have a simpler 
architecture than the spleen and lymph nodes in that they consist of a dense 
cluster of lymphocytes without a surrounding fibrous capsule. These 
nodules are located in the respiratory and digestive tracts, areas routinely 
exposed to environmental pathogens. 


Tonsils are lymphoid nodules located along the inner surface of the pharynx 
and are important in developing immunity to oral pathogens ([link]). The 
tonsil located at the back of the throat, the pharyngeal tonsil, is sometimes 
referred to as the adenoid when swollen. Such swelling is an indication of 
an active immune response to infection. Histologically, tonsils do not 
contain a complete capsule, and the epithelial layer invaginates deeply into 
the interior of the tonsil to form tonsillar crypts. These structures, which 
accumulate all sorts of materials taken into the body through eating and 
breathing, actually “encourage” pathogens to penetrate deep into the 


tonsillar tissues where they are acted upon by numerous lymphoid follicles 
and eliminated. This seems to be the major function of tonsils—to help 
children’s bodies recognize, destroy, and develop immunity to common 
environmental pathogens so that they will be protected in their later lives. 
Tonsils are often removed in those children who have recurring throat 
infections, especially those involving the palatine tonsils on either side of 
the throat, whose swelling may interfere with their breathing and/or 
swallowing. 

Locations and Histology of the Tonsils 


(a) Locations of the tonsils 


Brain 


Palatine Sphenoidal sinus 


tonsil 


Palatine Sphenoid bone 
bone 
Tongue ie in 
Mandible Nasopharynx 
Hyoid 
Trachea 
Esophagus 
Palatine Hard palate 
tonsil 
Soft palate 
Uvula 


Lingual 
tonsil Palatine tonsils 


(swollen due to infection) 
Epiglottis 


Tongue 


(b) Histology of palatine tonsil 


Crypt 


Stratified 
squamous 
epithelium 


Germinal 
centers 


(a) The pharyngeal tonsil is located on the roof of the 
posterior superior wall of the nasopharynx. The 
palatine tonsils lay on each side of the pharynx. (b) A 
micrograph shows the palatine tonsil tissue. LM x 40. 
(Micrograph provided by the Regents of the 
University of Michigan Medical School © 2012) 


Note: 


— 
mess OPenstax COLLEGE 
Poo 

- 


fi 


ae 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. 


Mucosa-associated lymphoid tissue (MALT) consists of an aggregate of 
lymphoid follicles directly associated with the mucous membrane epithelia. 
MALT makes up dome-shaped structures found underlying the mucosa of 
the gastrointestinal tract, breast tissue, lungs, and eyes. Peyer’s patches, a 
type of MALT in the small intestine, are especially important for immune 
responses against ingested substances ({link]). Peyer’s patches contain 
specialized endothelial cells called M (or microfold) cells that sample 
material from the intestinal lumen and transport it to nearby follicles so that 
adaptive immune responses to potential pathogens can be mounted. 
Mucosa-associated Lymphoid Tissue (MALT) Nodule 


Peyer's patches 


LM ~x 40. (Micrograph provided by the Regents of the 
University of Michigan Medical School © 2012) 


Bronchus-associated lymphoid tissue (BALT) consists of lymphoid 
follicular structures with an overlying epithelial layer found along the 
bifurcations of the bronchi, and between bronchi and arteries. They also 
have the typically less-organized structure of other lymphoid nodules. 
These tissues, in addition to the tonsils, are effective against inhaled 
pathogens. 


Chapter Review 


The lymphatic system is a series of vessels, ducts, and trunks that remove 
interstitial fluid from the tissues and return it the blood. The lymphatics are 
also used to transport dietary lipids and cells of the immune system. Cells 
of the immune system all come from the hematopoietic system of the bone 
marrow. Primary lymphoid organs, the bone marrow and thymus gland, are 
the locations where lymphocytes of the adaptive immune system proliferate 
and mature. Secondary lymphoid organs are site in which mature 


lymphocytes congregate to mount immune responses. Many immune 
system cells use the lymphatic and circulatory systems for transport 
throughout the body to search for and then protect against pathogens. 


Interactive Link Questions 


Exercise: 


Problem: 


Visit this website for an overview of the lymphatic system. What are 
the three main components of the lymphatic system? 


Solution: 
The three main components are the lymph vessels, the lymph nodes, 
and the lymph. 
Exercise: 
Problem: 
Visit this website to learn about the many different cell types in the 


immune system and their very specialized jobs. What is the role of the 
dendritic cell in infection by HIV? 


Solution: 


The dendritic cell transports the virus to a lymph node. 


Review Questions 


Exercise: 


Problem: Which of the following cells is phagocytic? 


a. plasma cell 
b. macrophage 


c. B cell 
d. NK cell 


Solution: 


B 
Exercise: 
Problem: 


Which structure allows lymph from the lower right limb to enter the 
bloodstream? 


a. thoracic duct 

b. right lymphatic duct 
c. right lymphatic trunk 
d. left lymphatic trunk 


Solution: 


A 
Exercise: 


Problem: 


Which of the following cells is important in the innate immune 
response? 


a. B cells 

b. T cells 

c. macrophages 
d. plasma cells 


Solution: 


c 


Exercise: 


Problem: 


Which of the following cells would be most active in early, antiviral 
immune responses the first time one is exposed to pathogen? 


a. macrophage 

b. T cell 

c. neutrophil 

d. natural killer cell 


Solution: 


D 
Exercise: 


Problem: 


Which of the lymphoid nodules is most likely to see food antigens 
first? 


a. tonsils 

b. Peyer’s patches 

c. bronchus-associated lymphoid tissue 
d. mucosa-associated lymphoid tissue 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: 


Describe the flow of lymph from its origins in interstitial fluid to its 
emptying into the venous bloodstream. 


Solution: 


The lymph enters through lymphatic capillaries, and then into larger 
lymphatic vessels. The lymph can only go in one direction due to 
valves in the vessels. The larger lymphatics merge to form trunks that 
enter into the blood via lymphatic ducts. 


Glossary 


adaptive immune response 
relatively slow but very specific and effective immune response 
controlled by lymphocytes 


afferent lymphatic vessels 
lead into a lymph node 


antibody 
antigen-specific protein secreted by plasma cells; immunoglobulin 


antigen 
molecule recognized by the receptors of B and T lymphocytes 


barrier defenses 
antipathogen defenses deriving from a barrier that physically prevents 
pathogens from entering the body to establish an infection 


B cells 
lymphocytes that act by differentiating into an antibody-secreting 
plasma cell 


bone marrow 


tissue found inside bones; the site of all blood cell differentiation and 
maturation of B lymphocytes 


bronchus-associated lymphoid tissue (BALT) 
lymphoid nodule associated with the respiratory tract 


chyle 
lipid-rich lymph inside the lymphatic capillaries of the small intestine 


cisterna chyli 
bag-like vessel that forms the beginning of the thoracic duct 


efferent lymphatic vessels 
lead out of a lymph node 


germinal centers 
clusters of rapidly proliferating B cells found in secondary lymphoid 
tissues 


high endothelial venules 
vessels containing unique endothelial cells specialized to allow 
migration of lymphocytes from the blood to the lymph node 


immune system 
series of barriers, cells, and soluble mediators that combine to response 
to infections of the body with pathogenic organisms 


innate immune response 
rapid but relatively nonspecific immune response 


lymph 
fluid contained within the lymphatic system 


lymph node 
one of the bean-shaped organs found associated with the lymphatic 
vessels 


lymphatic capillaries 
smallest of the lymphatic vessels and the origin of lymph flow 


lymphatic system 
network of lymphatic vessels, lymph nodes, and ducts that carries 
lymph from the tissues and back to the bloodstream. 


lymphatic trunks 
large lymphatics that collect lymph from smaller lymphatic vessels and 
empties into the blood via lymphatic ducts 


lymphocytes 
white blood cells characterized by a large nucleus and small rim of 
cytoplasm 


lymphoid nodules 
unencapsulated patches of lymphoid tissue found throughout the body 


mucosa-associated lymphoid tissue (MALT) 
lymphoid nodule associated with the mucosa 


naive lymphocyte 
mature B or T cell that has not yet encountered antigen for the first 
time 


natural killer cell (NK) 
cytotoxic lymphocyte of innate immune response 


plasma cell 
differentiated B cell that is actively secreting antibody 


primary lymphoid organ 
site where lymphocytes mature and proliferate; red bone marrow and 
thymus gland 


right lymphatic duct 
drains lymph fluid from the upper right side of body into the right 
subclavian vein 


secondary lymphoid organs 


sites where lymphocytes mount adaptive immune responses; examples 
include lymph nodes and spleen 


spleen 
secondary lymphoid organ that filters pathogens from the blood (white 
pulp) and removes degenerating or damaged blood cells (red pulp) 


T cell 
lymphocyte that acts by secreting molecules that regulate the immune 
system or by causing the destruction of foreign cells, viruses, and 
cancer cells 


thoracic duct 
large duct that drains lymph from the lower limbs, left thorax, left 
upper limb, and the left side of the head 


thymocyte 
immature T cell found in the thymus 


thymus 
primary lymphoid organ; where T lymphocytes proliferate and mature 


tonsils 
lymphoid nodules associated with the nasopharynx 


Gross Anatomy of Urine Transport 
By the end of this section, you will be able to: 


e Identify the ureters, urinary bladder, and urethra, as well as their 
location, structure, histology, and function 

e Compare and contrast male and female urethras 

e Describe the micturition reflex 

e Describe voluntary and involuntary neural control of micturition 


Rather than start with urine formation, this section will start with urine 
excretion. Urine is a fluid of variable composition that requires specialized 
structures to remove it from the body safely and efficiently. Blood is 
filtered, and the filtrate is transformed into urine at a relatively constant rate 
throughout the day. This processed liquid is stored until a convenient time 
for excretion. All structures involved in the transport and storage of the 
urine are large enough to be visible to the naked eye. This transport and 
storage system not only stores the waste, but it protects the tissues from 
damage due to the wide range of pH and osmolarity of the urine, prevents 
infection by foreign organisms, and for the male, provides reproductive 
functions. 


Urethra 


The urethra transports urine from the bladder to the outside of the body for 
disposal. The urethra is the only urologic organ that shows any significant 
anatomic difference between males and females; all other urine transport 
structures are identical ([link]). 

Female and Male Urethras 


The urethra transports urine from the bladder to the outside of the 
body. This image shows (a) a female urethra and (b) a male urethra. 


Urinary bladder 


Pubic bone Ureter 


Ureter 
Seminal 
vesicle 


Uterus —~_ Ductus 
Urinary \A nN ~ 

bladder a . 
Pubic bone 


sas Feu bi | i 
Clitoris ee 7 Vagina tactig ji: 


Prostate 
gland 


Rectum 


Anus 


The urethra in both males and females begins inferior and central to the two 
ureteral openings forming the three points of a triangular-shaped area at the 
base of the bladder called the trigone (Greek tri- = “triangle” and the root 
of the word “trigonometry”). The urethra tracks posterior and inferior to the 
pubic symphysis (see [link ]a). In both males and females, the proximal 
urethra is lined by transitional epithelium, whereas the terminal portion is a 
nonkeratinized, stratified squamous epithelium. In the male, 
pseudostratified columnar epithelium lines the urethra between these two 
cell types. Voiding is regulated by an involuntary autonomic nervous 
system-controlled internal urinary sphincter, consisting of smooth muscle 
and voluntary skeletal muscle that forms the external urinary sphincter 
below it. 


Female Urethra 


The external urethral orifice is embedded in the anterior vaginal wall 
inferior to the clitoris, superior to the vaginal opening (introitus), and 
medial to the labia minora. Its short length, about 4 cm, is less of a barrier 
to fecal bacteria than the longer male urethra and the best explanation for 
the greater incidence of UTI in women. Voluntary control of the external 
urethral sphincter is a function of the pudendal nerve. It arises in the sacral 
region of the spinal cord, traveling via the S2—S4 nerves of the sacral 
plexus. 


Male Urethra 


The male urethra passes through the prostate gland immediately inferior to 
the bladder before passing below the pubic symphysis (see [link]b). The 
length of the male urethra varies between men but averages 20 cm in 
length. It is divided into four regions: the preprostatic urethra, the prostatic 
urethra, the membranous urethra, and the spongy or penile urethra. The 
preprostatic urethra is very short and incorporated into the bladder wall. 
The prostatic urethra passes through the prostate gland. During sexual 
intercourse, it receives sperm via the ejaculatory ducts and secretions from 
the seminal vesicles. Paired Cowper’s glands (bulbourethral glands) 
produce and secrete mucus into the urethra to buffer urethral pH during 
sexual stimulation. The mucus neutralizes the usually acidic environment 
and lubricates the urethra, decreasing the resistance to ejaculation. The 
membranous urethra passes through the deep muscles of the perineum, 
where it is invested by the overlying urethral sphincters. The spongy urethra 
exits at the tip (external urethral orifice) of the penis after passing through 
the corpus spongiosum. Mucous glands are found along much of the length 
of the urethra and protect the urethra from extremes of urine pH. 
Innervation is the same in both males and females. 


Bladder 


The urinary bladder collects urine from both ureters ({link]). The bladder 
lies anterior to the uterus in females, posterior to the pubic bone and 
anterior to the rectum. During late pregnancy, its capacity is reduced due to 
compression by the enlarging uterus, resulting in increased frequency of 
urination. In males, the anatomy is similar, minus the uterus, and with the 
addition of the prostate inferior to the bladder. The bladder is partially 
retroperitoneal (outside the peritoneal cavity) with its peritoneal-covered 
“dome” projecting into the abdomen when the bladder is distended with 
urine. 

Bladder 


Ureter 


Peritoneum 


Detrusor 
muscle 


Ureteral 


, Transitional epithelium 
openings 


Lamina propria 
Submucosa 


Internal urethral 


sphincter | 
External urethral =a 
sphincter 


(a) (b) 


(a) Anterior cross section of the bladder. (b) The detrusor 
muscle of the bladder (source: monkey tissue) LM x 
448. (Micrograph provided by the Regents of the 
University of Michigan Medical School © 2012) 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. 


The bladder is a highly distensible organ comprised of irregular 
crisscrossing bands of smooth muscle collectively called the detrusor 
muscle. The interior surface is made of transitional cellular epithelium that 
is structurally suited for the large volume fluctuations of the bladder. When 
empty, it resembles columnar epithelia, but when stretched, it “transitions” 
(hence the name) to a squamous appearance (see [link]). Volumes in adults 
can range from nearly zero to 500-600 mL. 


The detrusor muscle contracts with significant force in the young. The 
bladder’s strength diminishes with age, but voluntary contractions of 
abdominal skeletal muscles can increase intra-abdominal pressure to 
promote more forceful bladder emptying. Such voluntary contraction is also 
used in forceful defecation and childbirth. 


Micturition Reflex 


Micturition is a less-often used, but proper term for urination or voiding. It 
results from an interplay of involuntary and voluntary actions by the 
internal and external urethral sphincters. When bladder volume reaches 
about 150 mL, an urge to void is sensed but is easily overridden. Voluntary 
control of urination relies on consciously preventing relaxation of the 
external urethral sphincter to maintain urinary continence. As the bladder 
fills, subsequent urges become harder to ignore. Ultimately, voluntary 
constraint fails with resulting incontinence, which will occur as bladder 
volume approaches 300 to 400 mL. 


Normal micturition is a result of stretch receptors in the bladder wall that 
transmit nerve impulses to the sacral region of the spinal cord to generate a 
spinal reflex. The resulting parasympathetic neural outflow causes 
contraction of the detrusor muscle and relaxation of the involuntary internal 
urethral sphincter. At the same time, the spinal cord inhibits somatic motor 
neurons, resulting in the relaxation of the skeletal muscle of the external 
urethral sphincter. The micturition reflex is active in infants but with 
maturity, children learn to override the reflex by asserting external sphincter 
control, thereby delaying voiding (potty training). This reflex may be 
preserved even in the face of spinal cord injury that results in paraplegia or 


quadriplegia. However, relaxation of the external sphincter may not be 
possible in all cases, and therefore, periodic catheterization may be 
necessary for bladder emptying. 


Nerves involved in the control of urination include the hypogastric, pelvic, 
and pudendal ([link]). Voluntary micturition requires an intact spinal cord 
and functional pudendal nerve arising from the sacral micturition center. 
Since the external urinary sphincter is voluntary skeletal muscle, actions by 
cholinergic neurons maintain contraction (and thereby continence) during 
filling of the bladder. At the same time, sympathetic nervous activity via the 
hypogastric nerves suppresses contraction of the detrusor muscle. With 
further bladder stretch, afferent signals traveling over sacral pelvic nerves 
activate parasympathetic neurons. This activates efferent neurons to release 
acetylcholine at the neuromuscular junctions, producing detrusor 
contraction and bladder emptying. 

Nerves Innervating the Urinary System 


Sacrum 


Uterus 


Urinary bladder 


Pubic bone 
Sphincter 
Clitoris 


Pudendal 
nerve 


Labium minora Anus 


Ureters 


The kidneys and ureters are completely retroperitoneal, and the bladder has 
a peritoneal covering only over the dome. As urine is formed, it drains into 
the calyces of the kidney, which merge to form the funnel-shaped renal 
pelvis in the hilum of each kidney. The renal pelvis narrows to become the 


ureter of each kidney. As urine passes through the ureter, it does not 
passively drain into the bladder but rather is propelled by waves of 
peristalsis. As the ureters enter the pelvis, they sweep laterally, hugging the 
pelvic walls. As they approach the bladder, they turn medially and pierce 
the bladder wall obliquely. This is important because it creates an one-way 
valve (a physiological sphincter rather than an anatomical sphincter) that 
allows urine into the bladder but prevents reflux of urine from the bladder 
back into the ureter. Children born lacking this oblique course of the ureter 
through the bladder wall are susceptible to “vesicoureteral reflux,” which 
dramatically increases their risk of serious UTI. Pregnancy also increases 
the likelihood of reflux and UTI. 


The ureters are approximately 30 cm long. The inner mucosa is lined with 
transitional epithelium ([link]) and scattered goblet cells that secrete 
protective mucus. The muscular layer of the ureter consists of longitudinal 
and circular smooth muscles that create the peristaltic contractions to move 
the urine into the bladder without the aid of gravity. Finally, a loose 
adventitial layer composed of collagen and fat anchors the ureters between 
the parietal peritoneum and the posterior abdominal wall. 

Ureter 


Peristaltic contractions help to move urine 


through the lumen with contributions from fluid 
pressure and gravity. LM x 128. (Micrograph 
provided by the Regents of the University of 
Michigan Medical School © 2012) 


Chapter Review 


The urethra is the only urinary structure that differs significantly between 
males and females. This is due to the dual role of the male urethra in 
transporting both urine and semen. The urethra arises from the trigone area 
at the base of the bladder. Urination is controlled by an involuntary internal 
sphincter of smooth muscle and a voluntary external sphincter of skeletal 
muscle. The shorter female urethra contributes to the higher incidence of 
bladder infections in females. The male urethra receives secretions from the 
prostate gland, Cowper’s gland, and seminal vesicles as well as sperm. The 
bladder is largely retroperitoneal and can hold up to 500-600 mL urine. 
Micturition is the process of voiding the urine and involves both 
involuntary and voluntary actions. Voluntary control of micturition requires 
a mature and intact sacral micturition center. It also requires an intact spinal 
cord. Loss of control of micturition is called incontinence and results in 
voiding when the bladder contains about 250 mL urine. The ureters are 
retroperitoneal and lead from the renal pelvis of the kidney to the trigone 
area at the base of the bladder. A thick muscular wall consisting of 
longitudinal and circular smooth muscle helps move urine toward the 
bladder by way of peristaltic contractions. 


Review Questions 


Exercise: 


Problem: Peristaltic contractions occur in the 


a. urethra 
b. bladder 


c. ureters 
d. urethra, bladder, and ureters 


Solution: 


C 
Exercise: 


Problem: 


Somatic motor neurons must be to relax the external 
urethral sphincter to allow urination. 


a. stimulated 
b. inhibited 


Solution: 


B 
Exercise: 


Problem: 
Which part of the urinary system is not completely retroperitoneal? 


a. kidneys 
b. ureters 

c. bladder 
d. nephrons 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: 


Why are females more likely to contract bladder infections than 
males? 


Solution: 


The longer urethra of males means bacteria must travel farther to the 
bladder to cause an infection. 


Exercise: 


Problem: Describe how forceful urination is accomplished. 


Solution: 


Forceful urination is accomplished by contraction of abdominal 
muscles. 


Glossary 


anatomical sphincter 
smooth or skeletal muscle surrounding the lumen of a vessel or hollow 
organ that can restrict flow when contracted 


detrusor muscle 
smooth muscle in the bladder wall; fibers run in all directions to 
reduce the size of the organ when emptying it of urine 


external urinary sphincter 
skeletal muscle; must be relaxed consciously to void urine 


internal urinary sphincter 
smooth muscle at the juncture of the bladder and urethra; relaxes as the 
bladder fills to allow urine into the urethra 


incontinence 
loss of ability to control micturition 


micturition 
also called urination or voiding 


physiological sphincter 
sphincter consisting of circular smooth muscle indistinguishable from 
adjacent muscle but possessing differential innervations, permitting its 
function as a sphincter; structurally weak 


retroperitoneal 
outside the peritoneal cavity; in the case of the kidney and ureters, 
between the parietal peritoneum and the abdominal wall 


sacral micturition center 
group of neurons in the sacral region of the spinal cord that controls 
urination; acts reflexively unless its action is modified by higher brain 
centers to allow voluntary urination 


trigone 
area at the base of the bladder marked by the two ureters in the 
posterior—lateral aspect and the urethral orifice in the anterior aspect 
oriented like points on a triangle 


urethra 
transports urine from the bladder to the outside environment 


Gross Anatomy of the Kidney 
By the end of this section, you will be able to: 


¢ Describe the external structure of the kidney, including its location, 
support structures, and covering 

¢ Identify the major internal divisions and structures of the kidney 

¢ Identify the major blood vessels associated with the kidney and trace 
the path of blood through the kidney 

e Compare and contrast the cortical and juxtamedullary nephrons 

e Name structures found in the cortex and medulla 

e Describe the physiological characteristics of the cortex and medulla 


The kidneys lie on either side of the spine in the retroperitoneal space 
between the parietal peritoneum and the posterior abdominal wall, well 
protected by muscle, fat, and ribs. They are roughly the size of your fist, 
and the male kidney is typically a bit larger than the female kidney. The 


kidneys are well vascularized, receiving about 25 percent of the cardiac 
output at rest. 


Note: 


— 
mess OPenstax COLLEGE 


There have never been sufficient kidney donations to provide a kidney to 
each person needing one. Watch this video to learn about the TED 
(Technology, Entertainment, Design) Conference held in March 2011. In 
this video, Dr. Anthony Atala discusses a cutting-edge technique in which 
anew kidney is “printed.” The successful utilization of this technology is 
still several years in the future, but imagine a time when you can print a 
replacement organ or tissue on demand. 


External Anatomy 


The left kidney is located at about the T12 to L3 vertebrae, whereas the 
right is lower due to slight displacement by the liver. Upper portions of the 
kidneys are somewhat protected by the eleventh and twelfth ribs ((link]). 
Each kidney weighs about 125-175 g in males and 115-155 g in females. 
They are about 11—14 cm in length, 6 cm wide, and 4 cm thick, and are 
directly covered by a fibrous capsule composed of dense, irregular 
connective tissue that helps to hold their shape and protect them. This 
capsule is covered by a shock-absorbing layer of adipose tissue called the 
renal fat pad, which in turn is encompassed by a tough renal fascia. The 
fascia and, to a lesser extent, the overlying peritoneum serve to firmly 
anchor the kidneys to the posterior abdominal wall in a retroperitoneal 
position. 

Kidneys 


Liver 


Kidney 
12th rib 


Ureter 


The kidneys are slightly protected by the ribs 
and are surrounded by fat for protection (not 
shown). 


On the superior aspect of each kidney is the adrenal gland. The adrenal 
cortex directly influences renal function through the production of the 
hormone aldosterone to stimulate sodium reabsorption. 


Internal Anatomy 


A frontal section through the kidney reveals an outer region called the renal 
cortex and an inner region called the medulla (({link]). The renal columns 
are connective tissue extensions that radiate downward from the cortex 
through the medulla to separate the most characteristic features of the 
medulla, the renal pyramids and renal papillae. The papillae are bundles 
of collecting ducts that transport urine made by nephrons to the calyces of 
the kidney for excretion. The renal columns also serve to divide the kidney 
into 6-8 lobes and provide a supportive framework for vessels that enter 
and exit the cortex. The pyramids and renal columns taken together 
constitute the kidney lobes. 

Left Kidney 


Cortical ——_ jie 


blood vessels _ Arcuate 


y) blood vessels 
| Interlobar ——_—_ ane 
| blood vessels 


Renal vein i) —_—— Major calyx 


Minor calyx 


aanel Renal pelvis 
Renal ena 


hilum nerve Pyramid 


Renal artery 
Papilla 
Medulla 


Renal column 
Ureter 


rt 
Capsule saute 


Renal Hilum 


The renal hilum is the entry and exit site for structures servicing the 
kidneys: vessels, nerves, lymphatics, and ureters. The medial-facing hila are 
tucked into the sweeping convex outline of the cortex. Emerging from the 
hilum is the renal pelvis, which is formed from the major and minor calyxes 


in the kidney. The smooth muscle in the renal pelvis funnels urine via 
peristalsis into the ureter. The renal arteries form directly from the 
descending aorta, whereas the renal veins return cleansed blood directly to 
the inferior vena cava. The artery, vein, and renal pelvis are arranged in an 
anterior-to-posterior order. 


Nephrons and Vessels 


The renal artery first divides into segmental arteries, followed by further 
branching to form interlobar arteries that pass through the renal columns to 
reach the cortex ([link]). The interlobar arteries, in turn, branch into arcuate 
arteries, cortical radiate arteries, and then into afferent arterioles. The 
afferent arterioles service about 1.3 million nephrons in each kidney. 

Blood Flow in the Kidney 


Peritubular capillaries 


Efferent 
arteriole 


Glomerulus 
Afferent 
arteriole 
Cortical 
radiate artery 


Arcuate 
artery 


Ee ~ 


Interlobar 
artery 


Segmental 
Interlobar 
vein 
Renal 
artery 


—— Renal vein 


Nephrons are the “functional units” of the kidney; they cleanse the blood 
and balance the constituents of the circulation. The afferent arterioles form 
a tuft of high-pressure capillaries about 200 pm in diameter, the 
glomerulus. The rest of the nephron consists of a continuous sophisticated 


tubule whose proximal end surrounds the glomerulus in an intimate 
embrace—this is Bowman’s capsule. The glomerulus and Bowman’s 
capsule together form the renal corpuscle. As mentioned earlier, these 
glomerular capillaries filter the blood based on particle size. After passing 
through the renal corpuscle, the capillaries form a second arteriole, the 
efferent arteriole ({link]). These will next form a capillary network around 
the more distal portions of the nephron tubule, the peritubular capillaries 
and vasa recta, before returning to the venous system. As the glomerular 
filtrate progresses through the nephron, these capillary networks recover 
most of the solutes and water, and return them to the circulation. Since a 
capillary bed (the glomerulus) drains into a vessel that in turn forms a 
second capillary bed, the definition of a portal system is met. This is the 
only portal system in which an arteriole is found between the first and 
second capillary beds. (Portal systems also link the hypothalamus to the 
anterior pituitary, and the blood vessels of the digestive viscera to the liver.) 
Blood Flow in the Nephron 


Glomerular capsule 


Efferent 
arteriole 
Afferent 
arteriole 


Proximal 
convoluted tubule 


Interlobular 
artery 


el 


Loop of 
A, the nephron 


Peritubular 
capillary 
network 


Urine flows into renal papilla 


The two capillary beds are clearly shown 
in this figure. The efferent arteriole is the 
connecting vessel between the glomerulus 
and the peritubular capillaries and vasa 
recta. 


Note: 


a] 


Visit this link to view an interactive tutorial of the flow of blood through 
the kidney. 


Cortex 


In a dissected kidney, it is easy to identify the cortex; it appears lighter in 
color compared to the rest of the kidney. All of the renal corpuscles as well 
as both the proximal convoluted tubules (PCTs) and distal convoluted 
tubules are found here. Some nephrons have a short loop of Henle that 
does not dip beyond the cortex. These nephrons are called cortical 
nephrons. About 15 percent of nephrons have long loops of Henle that 
extend deep into the medulla and are called juxtamedullary nephrons. 


Chapter Review 


As noted previously, the structure of the kidney is divided into two principle 
regions—the peripheral rim of cortex and the central medulla. The two 
kidneys receive about 25 percent of cardiac output. They are protected in 
the retroperitoneal space by the renal fat pad and overlying ribs and muscle. 
Ureters, blood vessels, lymph vessels, and nerves enter and leave at the 
renal hilum. The renal arteries arise directly from the aorta, and the renal 
veins drain directly into the inferior vena cava. Kidney function is derived 
from the actions of about 1.3 million nephrons per kidney; these are the 
“functional units.” A capillary bed, the glomerulus, filters blood and the 
filtrate is captured by Bowman’s capsule. A portal system is formed when 
the blood flows through a second capillary bed surrounding the proximal 
and distal convoluted tubules and the loop of Henle. Most water and solutes 
are recovered by this second capillary bed. This filtrate is processed and 
finally gathered by collecting ducts that drain into the minor calyces, which 


merge to form major calyces; the filtrate then proceeds to the renal pelvis 
and finally the ureters. 


Review Questions 


Exercise: 


Problem: 


The renal pyramids are separated from each other by extensions of the 
renal cortex called 


a. renal medulla 
b. minor calyces 
c. medullary cortices 
d. renal columns 


Solution: 


D 
Exercise: 


Problem: 
The primary structure found within the medulla is the 


a. loop of Henle 
b. minor calyces 
c. portal system 
d. ureter 


Solution: 


A 


Exercise: 


Problem:The right kidney is slightly lower because 


a. it is displaced by the liver 

b. it is displace by the heart 

c. it is slightly smaller 

d. it needs protection of the lower ribs 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem: What anatomical structures provide protection to the kidney? 
Solution: 


Retroperitoneal anchoring, renal fat pads, and ribs provide protection 
to the kidney. 

Exercise: 
Problem: 


How does the renal portal system differ from the hypothalamo— 
hypophyseal and digestive portal systems? 


Solution: 


The renal portal system has an artery between the first and second 
capillary bed. The others have a vein. 


Exercise: 


Problem: Name the structures found in the renal hilum. 


Solution: 


The structures found in the renal hilum are arteries, veins, ureters, 
lymphatics, and nerves. 


Glossary 


Bowman’s capsule 
cup-shaped sack lined by a simple squamous epithelium (parietal 
surface) and specialized cells called podocytes (visceral surface) that 
participate in the filtration process; receives the filtrate which then 
passes on to the PCTs 


calyces 
cup-like structures receiving urine from the collecting ducts where it 
passes on to the renal pelvis and ureter 


cortical nephrons 
nephrons with loops of Henle that do not extend into the renal medulla 


distal convoluted tubules 
portions of the nephron distal to the loop of Henle that receive 
hyposmotic filtrate from the loop of Henle and empty into collecting 
ducts 


efferent arteriole 
arteriole carrying blood from the glomerulus to the capillary beds 
around the convoluted tubules and loop of Henle; portion of the portal 
system 


glomerulus 
tuft of capillaries surrounded by Bowman’s capsule; filters the blood 
based on size 


juxtamedullary nephrons 
nephrons adjacent to the border of the cortex and medulla with loops 
of Henle that extend into the renal medulla 


loop of Henle 
descending and ascending portions between the proximal and distal 
convoluted tubules; those of cortical nephrons do not extend into the 
medulla, whereas those of juxtamedullary nephrons do extend into the 
medulla 


nephrons 
functional units of the kidney that carry out all filtration and 
modification to produce urine; consist of renal corpuscles, proximal 
and distal convoluted tubules, and descending and ascending loops of 
Henle; drain into collecting ducts 


medulla 
inner region of kidney containing the renal pyramids 


peritubular capillaries 
second capillary bed of the renal portal system; surround the proximal 
and distal convoluted tubules; associated with the vasa recta 


proximal convoluted tubules (PCTs) 
tortuous tubules receiving filtrate from Bowman’s capsule; most active 
part of the nephron in reabsorption and secretion 


renal columns 
extensions of the renal cortex into the renal medulla; separates the 
renal pyramids; contains blood vessels and connective tissues 


renal corpuscle 
consists of the glomerulus and Bowman’s capsule 


renal cortex 
outer part of kidney containing all of the nephrons; some nephrons 
have loops of Henle extending into the medulla 


renal fat pad 
adipose tissue between the renal fascia and the renal capsule that 
provides protective cushioning to the kidney 


renal hilum 
recessed medial area of the kidney through which the renal artery, 
renal vein, ureters, lymphatics, and nerves pass 


renal papillae 
medullary area of the renal pyramids where collecting ducts empty 
urine into the minor calyces 


renal pyramids 
six to eight cone-shaped tissues in the medulla of the kidney 
containing collecting ducts and the loops of Henle of juxtamedullary 
nephrons 


vasa recta 
branches of the efferent arterioles that parallel the course of the loops 
of Henle and are continuous with the peritubular capillaries; with the 
glomerulus, form a portal system 


Microscopic Anatomy of the Kidney 
By the end of this section, you will be able to: 


e Distinguish the histological differences between the renal cortex and 
medulla 

e Describe the structure of the filtration membrane 

e Identify the major structures and subdivisions of the renal corpuscles, 
renal tubules, and renal capillaries 

e Discuss the function of the peritubular capillaries and vasa recta 

e Identify the location of the juxtaglomerular apparatus and describe the 
cells that line it 

e Describe the histology of the proximal convoluted tubule, loop of 
Henle, distal convoluted tubule, and collecting ducts 


The renal structures that conduct the essential work of the kidney cannot be 
seen by the naked eye. Only a light or electron microscope can reveal these 
structures. Even then, serial sections and computer reconstruction are 
necessary to give us a comprehensive view of the functional anatomy of the 
nephron and its associated blood vessels. 


Nephrons: The Functional Unit 


Nephrons take a simple filtrate of the blood and modify it into urine. Many 
changes take place in the different parts of the nephron before urine is 
created for disposal. The term forming urine will be used hereafter to 
describe the filtrate as it is modified into true urine. The principle task of 
the nephron population is to balance the plasma to homeostatic set points 
and excrete potential toxins in the urine. They do this by accomplishing 
three principle functions—filtration, reabsorption, and secretion. They also 
have additional secondary functions that exert control in three areas: blood 
pressure (via production of renin), red blood cell production (via the 
hormone EPO), and calcium absorption (via conversion of calcidiol into 
calcitriol, the active form of vitamin D). 


Renal Corpuscle 


As discussed earlier, the renal corpuscle consists of a tuft of capillaries 
called the glomerulus that is largely surrounded by Bowman’s (glomerular) 
capsule. The glomerulus is a high-pressure capillary bed between afferent 
and efferent arterioles. Bowman’s capsule surrounds the glomerulus to form 
a lumen, and captures and directs this filtrate to the PCT. The outermost 
part of Bowman’s capsule, the parietal layer, is a simple squamous 
epithelium. It transitions onto the glomerular capillaries in an intimate 
embrace to form the visceral layer of the capsule. Here, the cells are not 
squamous, but uniquely shaped cells (podocytes) extending finger-like 
arms (pedicels) to cover the glomerular capillaries ((link]). These 
projections interdigitate to form filtration slits, leaving small gaps between 
the digits to form a sieve. As blood passes through the glomerulus, 10 to 20 
percent of the plasma filters between these sieve-like fingers to be captured 
by Bowman’s capsule and funneled to the PCT. Where the fenestrae 
(windows) in the glomerular capillaries match the spaces between the 
podocyte “fingers,” the only thing separating the capillary lumen and the 
lumen of Bowman’s capsule is their shared basement membrane ([link]). 
These three features comprise what is known as the filtration membrane. 
This membrane permits very rapid movement of filtrate from capillary to 
capsule though pores that are only 70 nm in diameter. 

Podocytes 


Cell 
Filtration bodies 


slits 


Capillary 


(b) 


Podocytes interdigitate with structures called pedicels 

and filter substances in a way similar to fenestrations. 
In (a), the large cell body can be seen at the top right 
comer, with branches extending from the cell body. 
The smallest finger-like extensions are the pedicels. 


Pedicels on one podocyte always interdigitate with the 
pedicels of another podocyte. (b) This capillary has 
three podocytes wrapped around it. 


Fenestrated Capillary 


Basement lle ae 


Endothelium 


Fenestrations 


Fenestrations allow many substances 
to diffuse from the blood based 
primarily on size. 


The fenestrations prevent filtration of blood cells or large proteins, but 
allow most other constituents through. These substances cross readily if 
they are less than 4 nm in size and most pass freely up to 8 nm in size. An 
additional factor affecting the ability of substances to cross this barrier is 
their electric charge. The proteins associated with these pores are negatively 
charged, so they tend to repel negatively charged substances and allow 
positively charged substances to pass more readily. The basement 
membrane prevents filtration of medium-to-large proteins such as globulins. 
There are also mesangial cells in the filtration membrane that can contract 
to help regulate the rate of filtration of the glomerulus. Overall, filtration is 
regulated by fenestrations in capillary endothelial cells, podocytes with 
filtration slits, membrane charge, and the basement membrane between 


capillary cells. The result is the creation of a filtrate that does not contain 
cells or large proteins, and has a slight predominance of positively charged 
substances. 


Lying just outside Bowman’s capsule and the glomerulus is the 
juxtaglomerular apparatus (JGA) ([link]). At the juncture where the 
afferent and efferent arterioles enter and leave Bowman’s capsule, the initial 
part of the distal convoluted tubule (DCT) comes into direct contact with 
the arterioles. The wall of the DCT at that point forms a part of the JGA 
known as the macula densa. This cluster of cuboidal epithelial cells 
monitors the fluid composition of fluid flowing through the DCT. In 
response to the concentration of Na” in the fluid flowing past them, these 
cells release paracrine signals. They also have a single, nonmotile cilium 
that responds to the rate of fluid movement in the tubule. The paracrine 
signals released in response to changes in flow rate and Na* concentration 
are adenosine triphosphate (ATP) and adenosine. 

Juxtaglomerular Apparatus and Glomerulus 


(a) The JGA allows specialized cells to monitor the composition of the 
fluid in the DCT and adjust the glomerular filtration rate. (b) This 
micrograph shows the glomerulus and surrounding structures. LM x 
1540. (Micrograph provided by the Regents of University of Michigan 
Medical School © 2012) 


Podocyte ; Macula densa 


Proximal 
convoluted 
tubule 


Brush 
border 


Juxtaglomerular 


Distal 
cells 


Proximal convoluted 
convoluted tubule 


Renal 
tubule 


nerve Basement 


membrane 


(a) (b) 


Afferent arteriole 


A second cell type in this apparatus is the juxtaglomerular cell. This is a 
modified, smooth muscle cell lining the afferent arteriole that can contract 
or relax in response to ATP or adenosine released by the macula densa. 
Such contraction and relaxation regulate blood flow to the glomerulus. If 


the osmolarity of the filtrate is too high (hyperosmotic), the juxtaglomerular 
cells will contract, decreasing the glomerular filtration rate (GFR) so less 
plasma is filtered, leading to less urine formation and greater retention of 
fluid. This will ultimately decrease blood osmolarity toward the physiologic 
norm. If the osmolarity of the filtrate is too low, the juxtaglomerular cells 
will relax, increasing the GFR and enhancing the loss of water to the urine, 
causing blood osmolarity to rise. In other words, when osmolarity goes up, 
filtration and urine formation decrease and water is retained. When 
osmolarity goes down, filtration and urine formation increase and water is 
lost by way of the urine. The net result of these opposing actions is to keep 
the rate of filtration relatively constant. A second function of the macula 
densa cells is to regulate renin release from the juxtaglomerular cells of the 
afferent arteriole ((link]). Active renin is a protein comprised of 304 amino 
acids that cleaves several amino acids from angiotensinogen to produce 
angiotensin I. Angiotensin I is not biologically active until converted to 
angiotensin II by angiotensin-converting enzyme (ACE) from the lungs. 
Angiotensin IT is a systemic vasoconstrictor that helps to regulate blood 
pressure by increasing it. Angiotensin II also stimulates the release of the 
steroid hormone aldosterone from the adrenal cortex. Aldosterone 
stimulates Na* reabsorption by the kidney, which also results in water 
retention and increased blood pressure. 

Conversion of Angiotensin I to Angiotensin II 


Macula densa 
senses low fluid 
flow or low Na* 
concentration 


Juxtaglomerular Angiotensin-converting 
cells secrete renin enzyme (ACE) in 
pulmonary blood 


Widespread vasoconstriction 


Kidney releases enzyme 
renin into blood 


Adrenal cortex 
to secrete 


Enzyme 


Angiotensin | 3 Aldosterone 
reaction 


ersten Angiotensin II Stimulates 


Liver releases 
angiotensinogen 
into blood 


ADH causes aquaporins to 
move to the collecting duct 
plasma membrane, which 

increases water reabsorption 


>| Aldosterone stimulates Nat 
uptake on the apical cell 
membrane in the distal 
convoluted tubule and 
collecting ducts 


The enzyme renin converts the pro-enzyme angiotensin I; the lung- 
derived enzyme ACE converts angiotensin I into active angiotensin 
i. 


Proximal Convoluted Tubule (PCT) 


Filtered fluid collected by Bowman’s capsule enters into the PCT. It is 
called convoluted due to its tortuous path. Simple cuboidal cells form this 
tubule with prominent microvilli on the luminal surface, forming a brush 
border. These microvilli create a large surface area to maximize the 
absorption and secretion of solutes (Na*, Cl’, glucose, etc.), the most 
essential function of this portion of the nephron. These cells actively 
transport ions across their membranes, so they possess a high concentration 
of mitochondria in order to produce sufficient ATP. 


Loop of Henle 


The descending and ascending portions of the loop of Henle (sometimes 
referred to as the nephron loop) are, of course, just continuations of the 
same tubule. They run adjacent and parallel to each other after having made 
a hairpin turn at the deepest point of their descent. The descending loop of 
Henle consists of an initial short, thick portion and long, thin portion, 
whereas the ascending loop consists of an initial short, thin portion 
followed by a long, thick portion. The descending thick portion consists of 
simple cuboidal epithelium similar to that of the PCT. The descending and 
ascending thin portions consists of simple squamous epithelium. As you 
will see later, these are important differences, since different portions of the 
loop have different permeabilities for solutes and water. The ascending 
thick portion consists of simple cuboidal epithelium similar to the DCT. 


Distal Convoluted Tubule (DCT) 


The DCT, like the PCT, is very tortuous and formed by simple cuboidal 
epithelium, but it is shorter than the PCT. These cells are not as active as 
those in the PCT; thus, there are fewer microvilli on the apical surface. 
However, these cells must also pump ions against their concentration 
gradient, so you will find of large numbers of mitochondria, although fewer 
than in the PCT. 


Collecting Ducts 


The collecting ducts are continuous with the nephron but not technically 
part of it. In fact, each duct collects filtrate from several nephrons for final 
modification. Collecting ducts merge as they descend deeper in the medulla 
to form about 30 terminal ducts, which empty at a papilla. They are lined 
with simple squamous epithelium with receptors for ADH. When 
stimulated by ADH, these cells will insert aquaporin channel proteins into 
their membranes, which as their name suggests, allow water to pass from 
the duct lumen through the cells and into the interstitial spaces to be 


recovered by the vasa recta. This process allows for the recovery of large 
amounts of water from the filtrate back into the blood. In the absence of 
ADH, these channels are not inserted, resulting in the excretion of water in 
the form of dilute urine. Most, if not all, cells of the body contain aquaporin 
molecules, whose channels are so small that only water can pass. At least 
10 types of aquaporins are known in humans, and six of those are found in 
the kidney. The function of all aquaporins is to allow the movement of 
water across the lipid-rich, hydrophobic cell membrane ([link]). 

Aquaporin Water Channel 


Water 
channel 


Cell membrane 


Positive charges inside the channel prevent 
the leakage of electrolytes across the cell 
membrane, while allowing water to move 

due to osmosis. 


Chapter Review 


The functional unit of the kidney, the nephron, consists of the renal 
corpuscle, PCT, loop of Henle, and DCT. Cortical nephrons have short 
loops of Henle, whereas juxtamedullary nephrons have long loops of Henle 
extending into the medulla. About 15 percent of nephrons are 
juxtamedullary. The glomerulus is a capillary bed that filters blood 
principally based on particle size. The filtrate is captured by Bowman’s 
capsule and directed to the PCT. A filtration membrane is formed by the 
fused basement membranes of the podocytes and the capillary endothelial 
cells that they embrace. Contractile mesangial cells further perform a role in 


regulating the rate at which the blood is filtered. Specialized cells in the 
JGA produce paracrine signals to regulate blood flow and filtration rates of 
the glomerulus. Other JGA cells produce the enzyme renin, which plays a 
central role in blood pressure regulation. The filtrate enters the PCT where 
absorption and secretion of several substances occur. The descending and 
ascending limbs of the loop of Henle consist of thick and thin segments. 
Absorption and secretion continue in the DCT but to a lesser extent than in 
the PCT. Each collecting duct collects forming urine from several nephrons 
and responds to the posterior pituitary hormone ADH by inserting 
aquaporin water channels into the cell membrane to fine tune water 
recovery. 


Review Questions 


Exercise: 


Problem:Blood filtrate is captured in the lumen of the 


a. glomerulus 

b. Bowman’s capsule 
c. calyces 

d. renal papillae 


Solution: 


B 
Exercise: 


Problem: 
What are the names of the capillaries following the efferent arteriole? 


a. arcuate and medullary 

b. interlobar and interlobular 
c. peritubular and vasa recta 
d. peritubular and medullary 


Solution: 
C 
Exercise: 
Problem:The functional unit of the kidney is called 


a. the renal hilus 

b. the renal corpuscle 
c. the nephron 

d. Bowman’s capsule 


Solution: 


C 


Critical Thinking Questions 


Exercise: 


Problem: Which structures make up the renal corpuscle? 


Solution: 


The structures that make up the renal corpuscle are the glomerulus, 
Bowman’s capsule, and PCT. 


Exercise: 


Problem: 


What are the major structures comprising the filtration membrane? 


Solution: 


The major structures comprising the filtration membrane are 
fenestrations and podocyte fenestra, fused basement membrane, and 
filtration slits. 


Glossary 


angiotensin-converting enzyme (ACE) 
enzyme produced by the lungs that catalyzes the reaction of inactive 
angiotensin I into active angiotensin II 


angiotensin I 
protein produced by the enzymatic action of renin on angiotensinogen; 
inactive precursor of angiotensin II 


angiotensin II 
protein produced by the enzymatic action of ACE on inactive 
angiotensin I; actively causes vasoconstriction and stimulates 
aldosterone release by the adrenal cortex 


angiotensinogen 
inactive protein in the circulation produced by the liver; precursor of 
angiotensin I; must be modified by the enzymes renin and ACE to be 
activated 


aquaporin 
protein-forming water channels through the lipid bilayer of the cell; 
allows water to cross; activation in the collecting ducts is under the 
control of ADH 


brush border 
formed by microvilli on the surface of certain cuboidal cells; in the 
kidney it is found in the PCT; increases surface area for absorption in 
the kidney 


fenestrations 
small windows through a cell, allowing rapid filtration based on size; 
formed in such a way as to allow substances to cross through a cell 


without mixing with cell contents 


filtration slits 
formed by pedicels of podocytes; substances filter between the 
pedicels based on size 


forming urine 
filtrate undergoing modifications through secretion and reabsorption 
before true urine is produced 


juxtaglomerular apparatus (JGA) 
located at the juncture of the DCT and the afferent and efferent 
arterioles of the glomerulus; plays a role in the regulation of renal 
blood flow and GFR 


juxtaglomerular cell 
modified smooth muscle cells of the afferent arteriole; secretes renin in 
response to a drop in blood pressure 


macula densa 
cells found in the part of the DCT forming the JGA; sense Na* 
concentration in the forming urine 


mesangial 
contractile cells found in the glomerulus; can contract or relax to 
regulate filtration rate 


pedicels 
finger-like projections of podocytes surrounding glomerular 
capillaries; interdigitate to form a filtration membrane 


podocytes 
cells forming finger-like processes; form the visceral layer of 
Bowman’s capsule; pedicels of the podocytes interdigitate to form a 
filtration membrane 


renin 


enzyme produced by juxtaglomerular cells in response to decreased 
blood pressure or sympathetic nervous activity; catalyzes the 
conversion of angiotensinogen into angiotensin I 


An Overview of the Endocrine System 
By the end of this section, you will be able to: 


e Distinguish the types of intercellular communication, their importance, 
mechanisms, and effects 

e Identify the major organs and tissues of the endocrine system and their 
location in the body 


Communication is a process in which a sender transmits signals to one or 
more receivers to control and coordinate actions. In the human body, two 
major organ systems participate in relatively “long distance” 
communication: the nervous system and the endocrine system. Together, 
these two systems are primarily responsible for maintaining homeostasis in 
the body. 


Neural and Endocrine Signaling 


The nervous system uses two types of intercellular communication— 
electrical and chemical signaling—either by the direct action of an 
electrical potential, or in the latter case, through the action of chemical 
neurotransmitters such as serotonin or norepinephrine. Neurotransmitters 
act locally and rapidly. When an electrical signal in the form of an action 
potential arrives at the synaptic terminal, they diffuse across the synaptic 
cleft (the gap between a sending neuron and a receiving neuron or muscle 
cell). Once the neurotransmitters interact (bind) with receptors on the 
receiving (post-synaptic) cell, the receptor stimulation is transduced into a 
response such as continued electrical signaling or modification of cellular 
response. The target cell responds within milliseconds of receiving the 
chemical “message”; this response then ceases very quickly once the neural 
signaling ends. In this way, neural communication enables body functions 
that involve quick, brief actions, such as movement, sensation, and 
cognition.In contrast, the endocrine system uses just one method of 
communication: chemical signaling. These signals are sent by the endocrine 
organs, which secrete chemicals—the hormone— into the extracellular 
fluid. Hormones are transported primarily via the bloodstream throughout 
the body, where they bind to receptors on target cells, inducing a 
characteristic response. As a result, endocrine signaling requires more time 


than neural signaling to prompt a response in target cells, though the precise 
amount of time varies with different hormones. For example, the hormones 
released when you are confronted with a dangerous or frightening situation, 
called the fight-or-flight response, occur by the release of adrenal hormones 
—epinephrine and norepinephrine—within seconds. In contrast, it may take 
up to 48 hours for target cells to respond to certain reproductive hormones. 


fli: fm) 

x Fr 
= 

openstax COLLEGE” 


1 


Visit this link to watch an animation of the events that occur when a 
hormone binds to a cell membrane receptor. What is the secondary 
messenger made by adenylyl cyclase during the activation of liver cells by 
epinephrine? 


In addition, endocrine signaling is typically less specific than neural 
signaling. The same hormone may play a role in a variety of different 
physiological processes depending on the target cells involved. For 
example, the hormone oxytocin promotes uterine contractions in women in 
labor. It is also important in breastfeeding, and may be involved in the 
sexual response and in feelings of emotional attachment in both males and 
females. 


In general, the nervous system involves quick responses to rapid changes in 
the external environment, and the endocrine system is usually slower acting 
—taking care of the internal environment of the body, maintaining 
homeostasis, and controlling reproduction ({link]). So how does the fight- 
or-flight response that was mentioned earlier happen so quickly if hormones 
are usually slower acting? It is because the two systems are connected. It is 


the fast action of the nervous system in response to the danger in the 
environment that stimulates the adrenal glands to secrete their hormones. 
As a result, the nervous system can cause rapid endocrine responses to keep 
up with sudden changes in both the external and internal environments 


when necessary. 


Endocrine and Nervous Systems 


Endocrine 
system 
Siena Chemical 
mechanism(s) 
Primary chemical 
: Hormones 
signal 
Distance traveled Long or short 
Response time Fast or slow 
Environment targeted Internal 


Structures of the Endocrine System 


Nervous system 


Chemical/electrical 


Neurotransmitters 


Always short 
Always fast 


Internal and 
external 


The endocrine system consists of cells, tissues, and organs that secrete 
hormones as a primary or secondary function. The endocrine gland is the 
major player in this system. The primary function of these ductless glands is 
to secrete their hormones directly into the surrounding fluid. The interstitial 
fluid and the blood vessels then transport the hormones throughout the 


body. The endocrine system includes the pituitary, thyroid, parathyroid, 
adrenal, and pineal glands ({link]). Some of these glands have both 
endocrine and non-endocrine functions. For example, the pancreas contains 
cells that function in digestion as well as cells that secrete the hormones 
insulin and glucagon, which regulate blood glucose levels. The 
hypothalamus, thymus, heart, kidneys, stomach, small intestine, liver, skin, 
female ovaries, and male testes are other organs that contain cells with 
endocrine function. Moreover, adipose tissue has long been known to 
produce hormones, and recent research has revealed that even bone tissue 
has endocrine functions. 

Endocrine System 


Pineal gland 


Thalamus 


Pituitary gland 


Thyroid cartilage 
of the larynx 


Thyroid gland 


Thymus Parathyroid glands 


(on posterior side 
of thyroid) 


Trachea 


Adrenal glands 


Pancreas 


Uterus 


Ovaries (female) 


Testes (male) 


Endocrine glands and cells are located throughout the body 
and play an important role in homeostasis. 


The ductless endocrine glands are not to be confused with the body’s 
exocrine system, whose glands release their secretions through ducts. 
Examples of exocrine glands include the sebaceous and sweat glands of the 
skin. As just noted, the pancreas also has an exocrine function: most of its 
cells secrete pancreatic juice through the pancreatic and accessory ducts to 
the lumen of the small intestine. 


Other Types of Chemical Signaling 


In endocrine signaling, hormones secreted into the extracellular fluid 
diffuse into the blood or lymph, and can then travel great distances 
throughout the body. In contrast, autocrine signaling takes place within the 
same cell. An autocrine (auto- = “self”) is a chemical that elicits a response 
in the same cell that secreted it. Interleukin-1, or IL-1, is a signaling 
molecule that plays an important role in inflammatory response. The cells 
that secrete IL-1 have receptors on their cell surface that bind these 
molecules, resulting in autocrine signaling. 


Local intercellular communication is the province of the paracrine, also 
called a paracrine factor, which is a chemical that induces a response in 
neighboring cells. Although paracrines may enter the bloodstream, their 
concentration is generally too low to elicit a response from distant tissues. 
A familiar example to those with asthma is histamine, a paracrine that is 
released by immune cells in the bronchial tree. Histamine causes the 
smooth muscle cells of the bronchi to constrict, narrowing the airways. 
Another example is the neurotransmitters of the nervous system, which act 
only locally within the synaptic cleft. 


Note: 

Career Connections 

Endocrinologist 

Endocrinology is a specialty in the field of medicine that focuses on the 
treatment of endocrine system disorders. Endocrinologists—medical 
doctors who specialize in this field—are experts in treating diseases 
associated with hormonal systems, ranging from thyroid disease to diabetes 


mellitus. Endocrine surgeons treat endocrine disease through the removal, 
or resection, of the affected endocrine gland. 

Patients who are referred to endocrinologists may have signs and 
symptoms or blood test results that suggest excessive or impaired 
functioning of an endocrine gland or endocrine cells. The endocrinologist 
may order additional blood tests to determine whether the patient’s 
hormonal levels are abnormal, or they may stimulate or suppress the 
function of the suspect endocrine gland and then have blood taken for 
analysis. Treatment varies according to the diagnosis. Some endocrine 
disorders, such as type 2 diabetes, may respond to lifestyle changes such as 
modest weight loss, adoption of a healthy diet, and regular physical 
activity. Other disorders may require medication, such as hormone 
replacement, and routine monitoring by the endocrinologist. These include 
disorders of the pituitary gland that can affect growth and disorders of the 
thyroid gland that can result in a variety of metabolic problems. 

Some patients experience health problems as a result of the normal decline 
in hormones that can accompany aging. These patients can consult with an 
endocrinologist to weigh the risks and benefits of hormone replacement 
therapy intended to boost their natural levels of reproductive hormones. 

In addition to treating patients, endocrinologists may be involved in 
research to improve the understanding of endocrine system disorders and 
develop new treatments for these diseases. 


Chapter Review 


The endocrine system consists of cells, tissues, and organs that secrete 
hormones critical to homeostasis. The body coordinates its functions 
through two major types of communication: neural and endocrine. Neural 
communication includes both electrical and chemical signaling between 
neurons and target cells. Endocrine communication involves chemical 
signaling via the release of hormones into the extracellular fluid. From 
there, hormones diffuse into the bloodstream and may travel to distant body 
regions, where they elicit a response in target cells. Endocrine glands are 
ductless glands that secrete hormones. Many organs of the body with other 


primary functions—such as the heart, stomach, and kidneys—also have 
hormone-secreting cells. 


Interactive Link Questions 


Exercise: 


Problem: 


Visit this link to watch an animation of the events that occur when a 
hormone binds to a cell membrane receptor. What is the secondary 
messenger made by adenylyl cyclase during the activation of liver 
cells by epinephrine? 


Solution: 


cAMP 


Review Questions 


Exercise: 


Problem:Endocrine glands 


a. secrete hormones that travel through a duct to the target organs 
b. release neurotransmitters into the synaptic cleft 

c. secrete chemical messengers that travel in the bloodstream 

d. include sebaceous glands and sweat glands 


Solution: 


G 


Exercise: 


Problem: 
Chemical signaling that affects neighboring cells is called 


a. autocrine 
b. paracrine 
c. endocrine 
d. neuron 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


Describe several main differences in the communication methods used 
by the endocrine system and the nervous system. 


Solution: 


The endocrine system uses chemical signals called hormones to 
convey information from one part of the body to a distant part of the 
body. Hormones are released from the endocrine cell into the 
extracellular environment, but then travel in the bloodstream to target 
tissues. This communication and response can take seconds to days. In 
contrast, neurons transmit electrical signals along their axons. At the 
axon terminal, the electrical signal prompts the release of a chemical 
signal called a neurotransmitter that carries the message across the 
synaptic cleft to elicit a response in the neighboring cell. This method 
of communication is nearly instantaneous, of very brief duration, and 
is highly specific. 


Exercise: 


Problem:Compare and contrast endocrine and exocrine glands. 


Solution: 


Endocrine glands are ductless. They release their secretion into the 
surrounding fluid, from which it enters the bloodstream or lymph to 
travel to distant cells. Moreover, the secretions of endocrine glands are 
hormones. Exocrine glands release their secretions through a duct that 
delivers the secretion to the target location. Moreover, the secretions of 
exocrine glands are not hormones, but compounds that have an 
immediate physiologic function. For example, pancreatic juice 
contains enzymes that help digest food. 


Exercise: 
Problem: 


True or false: Neurotransmitters are a special class of paracrines. 
Explain your answer. 


Solution: 


True. Neurotransmitters can be classified as paracrines because, upon 
their release from a neuron’s axon terminals, they travel across a 
microscopically small cleft to exert their effect on a nearby neuron or 
muscle cell. 


Glossary 


autocrine 
chemical signal that elicits a response in the same cell that secreted it 


endocrine gland 
tissue or organ that secretes hormones into the blood and lymph 
without ducts such that they may be transported to organs distant from 
the site of secretion 


endocrine system 
cells, tissues, and organs that secrete hormones as a primary or 
secondary function and play an integral role in normal bodily 
processes 


exocrine system 
cells, tissues, and organs that secrete substances directly to target 
tissues via glandular ducts 


hormone 
secretion of an endocrine organ that travels via the bloodstream or 
lymphatics to induce a response in target cells or tissues in another part 
of the body 


paracrine 
chemical signal that elicits a response in neighboring cells; also called 
paracrine factor 


The Pituitary Gland and Hypothalamus 
By the end of this section, you will be able to: 


e Explain the interrelationships of the anatomy and functions of the 
hypothalamus and the posterior and anterior lobes of the pituitary gland 

e Identify the two hormones released from the posterior pituitary, their 
target cells, and their principal actions 

e Identify the six hormones produced by the anterior lobe of the pituitary 
gland, their target cells, their principal actions, and their regulation by the 
hypothalamus 


The hypothalamus-—pituitary complex can be thought of as the “command 
center” of the endocrine system. This complex secretes several hormones that 
directly produce responses in target tissues, as well as hormones that regulate 
the synthesis and secretion of hormones of other glands. In addition, the 
hypothalamus-pituitary complex coordinates the messages of the endocrine 
and nervous systems. In many cases, a stimulus received by the nervous system 
must pass through the hypothalamus-pituitary complex to be translated into 
hormones that can initiate a response. 


The hypothalamus is a structure of the diencephalon of the brain located 
anterior and inferior to the thalamus ({link]). It has both neural and endocrine 
functions, producing and secreting many hormones. In addition, the 
hypothalamus is anatomically and functionally related to the pituitary gland 
(or hypophysis), a bean-sized organ suspended from it by a stem called the 
infundibulum (or pituitary stalk). The pituitary gland is cradled within the 
sellaturcica of the sphenoid bone of the skull. It consists of two lobes that arise 
from distinct parts of embryonic tissue: the posterior pituitary 
(neurohypophysis) is neural tissue, whereas the anterior pituitary (also known 
as the adenohypophysis) is glandular tissue that develops from the primitive 
digestive tract. The hormones secreted by the posterior and anterior pituitary, 
and the intermediate zone between the lobes are summarized in [link]. 
Hypothalamus—Pituitary Complex 


Thalamus 


Hypothalamus 


Infundibulum 
Anterior pituitary 


Posterior pituitary 


Se 
NE 


The hypothalamus region lies inferior and anterior to the 
thalamus. It connects to the pituitary gland by the stalk-like 
infundibulum. The pituitary gland consists of an anterior 
and posterior lobe, with each lobe secreting different 
hormones in response to signals from the hypothalamus. 


Pituitary Hormones 


Pituitary Associated Chemical 
lobe hormones class Effect 


Promotes 
Growth hormone 
Anterior (GH) Protein growth of 


body tissues 


Pituitary Hormones 


Pituitary 
lobe 


Anterior 


Anterior 


Anterior 


Anterior 


Anterior 


Posterior 


Associated 
hormones 


Prolactin (PRL) 


Thyroid-stimulating 
hormone (TSH) 


Adrenocorticotropic 
hormone (ACTH) 


Follicle-stimulating 
hormone (FSH) 


Luteinizing 
hormone (LH) 


Antidiuretic 
hormone (ADH) 


Chemical 
class 


Peptide 


Glycoprotein 


Peptide 


Glycoprotein 


Glycoprotein 


Peptide 


Effect 


Promotes 
milk 
production 
from 
mammary 
glands 


Stimulates 
thyroid 
hormone 
release from 
thyroid 


Stimulates 
hormone 
release by 
adrenal 
cortex 


Stimulates 
gamete 
production 
in gonads 


Stimulates 
androgen 

production 
by gonads 


Stimulates 
water 

reabsorption 
by kidneys 


Pituitary Hormones 


Pituitary Associated Chemical 
lobe hormones class 
Posterior Oxytocin Peptide 
Melanocyte- 
Intermediate 
stimulating Peptide 
zone 
hormone 


Posterior Pituitary 


Effect 


Stimulates 
uterine 
contractions 
during 
childbirth 


Stimulates 
melanin 
formation in 
melanocytes 


The posterior pituitary is actually an extension of the neurons of the 
paraventricular and supraoptic nuclei of the hypothalamus. The cell bodies of 
these regions rest in the hypothalamus, but their axons descend as the 
hypothalamic—hypophyseal tract within the infundibulum, and end in axon 


terminals that comprise the posterior pituitary ([Link]). 
Posterior Pituitary 


Neurosecretory cells of Neurosecretory cells of 
paraventricular nucleus supraoptic nucleus 


ADH release 


Hypothalamus 


Infundibulum 


Hypothalamohypophyseal 
tract 


Posterior 
pituitary 
Pituitary 


Anterior pituitary gland 


Capillary plexus 


Oitielease ADH release 


Neurosecretory cells in the hypothalamus release oxytocin 
(OT) or ADH into the posterior lobe of the pituitary gland. 
These hormones are stored or released into the blood via 
the capillary plexus. 


The posterior pituitary gland does not produce hormones, but rather stores and 
secretes hormones produced by the hypothalamus. The paraventricular nuclei 
produce the hormone oxytocin, whereas the supraoptic nuclei produce ADH. 
These hormones travel along the axons into storage sites in the axon terminals 
of the posterior pituitary. In response to signals from the same hypothalamic 
neurons, the hormones are released from the axon terminals into the 
bloodstream. 


Oxytocin 


When fetal development is complete, the peptide-derived hormone oxytocin 
(tocia- = “childbirth”) stimulates uterine contractions and dilation of the cervix. 
Throughout most of pregnancy, oxytocin hormone receptors are not expressed 
at high levels in the uterus. Toward the end of pregnancy, the synthesis of 
oxytocin receptors in the uterus increases, and the smooth muscle cells of the 
uterus become more sensitive to its effects. Oxytocin is continually released 
throughout childbirth through a positive feedback mechanism. As noted earlier, 
oxytocin prompts uterine contractions that push the fetal head toward the 
cervix. In response, cervical stretching stimulates additional oxytocin to be 
synthesized by the hypothalamus and released from the pituitary. This increases 
the intensity and effectiveness of uterine contractions and prompts additional 
dilation of the cervix. The feedback loop continues until birth. 


Although the mother’s high blood levels of oxytocin begin to decrease 
immediately following birth, oxytocin continues to play a role in maternal and 
newborn health. First, oxytocin is necessary for the milk ejection reflex 
(commonly referred to as “let-down”) in breastfeeding women. As the newborn 
begins suckling, sensory receptors in the nipples transmit signals to the 
hypothalamus. In response, oxytocin is secreted and released into the 
bloodstream. Within seconds, cells in the mother’s milk ducts contract, ejecting 
milk into the infant’s mouth. Secondly, in both males and females, oxytocin is 
thought to contribute to parent-newborn bonding, known as attachment. 
Oxytocin is also thought to be involved in feelings of love and closeness, as 
well as in the sexual response. 


Antidiuretic Hormone (ADH) 


The solute concentration of the blood, or blood osmolarity, may change in 

response to the consumption of certain foods and fluids, as well as in response 
to disease, injury, medications, or other factors. Blood osmolarity is constantly 
monitored by osmoreceptors—specialized cells within the hypothalamus that 
are particularly sensitive to the concentration of sodium ions and other solutes. 


In response to high blood osmolarity, which can occur during dehydration or 
following a very salty meal, the osmoreceptors signal the posterior pituitary to 
release antidiuretic hormone (ADH). The target cells of ADH are located in 
the tubular cells of the kidneys. Its effect is to increase epithelial permeability 
to water, allowing increased water reabsorption. The more water reabsorbed 


from the filtrate, the greater the amount of water that is returned to the blood 
and the less that is excreted in the urine. A greater concentration of water 
results in a reduced concentration of solutes. ADH is also known as vasopressin 
because, in very high concentrations, it causes constriction of blood vessels, 
which increases blood pressure by increasing peripheral resistance. The release 
of ADH is controlled by a negative feedback loop. As blood osmolarity 
decreases, the hypothalamic osmoreceptors sense the change and prompt a 
corresponding decrease in the secretion of ADH. As a result, less water is 
reabsorbed from the urine filtrate. 


Interestingly, drugs can affect the secretion of ADH. For example, alcohol 
consumption inhibits the release of ADH, resulting in increased urine 
production that can eventually lead to dehydration and a hangover. A disease 
called diabetes insipidus is characterized by chronic underproduction of ADH 
that causes chronic dehydration. Because little ADH is produced and secreted, 
not enough water is reabsorbed by the kidneys. Although patients feel thirsty, 
and increase their fluid consumption, this doesn’t effectively decrease the 
solute concentration in their blood because ADH levels are not high enough to 
trigger water reabsorption in the kidneys. Electrolyte imbalances can occur in 
severe cases of diabetes insipidus. 


Anterior Pituitary 


The anterior pituitary originates from the digestive tract in the embryo and 
migrates toward the brain during fetal development. There are three regions: 
the pars distalis is the most anterior, the pars intermedia is adjacent to the 
posterior pituitary, and the pars tuberalis is a slender “tube” that wraps the 
infundibulum. 


Recall that the posterior pituitary does not synthesize hormones, but merely 
stores them. In contrast, the anterior pituitary does manufacture hormones. 
However, the secretion of hormones from the anterior pituitary is regulated by 
two classes of hormones. These hormones—secreted by the hypothalamus—are 
the releasing hormones that stimulate the secretion of hormones from the 
anterior pituitary and the inhibiting hormones that inhibit secretion. 


Hypothalamic hormones are secreted by neurons, but enter the anterior 
pituitary through blood vessels ([{link]). Within the infundibulum is a bridge of 
capillaries that connects the hypothalamus to the anterior pituitary. This 


network, called the hypophyseal portal system, allows hypothalamic 
hormones to be transported to the anterior pituitary without first entering the 
systemic circulation. The system originates from the superior hypophyseal 
artery, which branches off the carotid arteries and transports blood to the 
hypothalamus. The branches of the superior hypophyseal artery form the 
hypophyseal portal system (see [link]). Hypothalamic releasing and inhibiting 
hormones travel through a primary capillary plexus to the portal veins, which 
carry them into the anterior pituitary. Hormones produced by the anterior 
pituitary (in response to releasing hormones) enter a secondary capillary 
plexus, and from there drain into the circulation. 

Anterior Pituitary 


@) Hypothalamus 
releases hormone 


Superior 
Hypothalamus hypophyseal 


Neurosecretory 
cells 


Infundibulum : WSs Primary capillary 
Hypophyseal y plexus of hypophyseal 
portal veins r portal system 


Posterior pituitary 


Anterior 


Pituitary gland 
Secondary capillary 
plexus of hypophyseal 
portal system 


@) Anterior pituitary @) Hypothalamus hormone stimulates 
hormone pituitary to release hormones 


The anterior pituitary manufactures seven hormones. The 
hypothalamus produces separate hormones that stimulate or 
inhibit hormone production in the anterior pituitary. 
Hormones from the hypothalamus reach the anterior 
pituitary via the hypophyseal portal system. 


The anterior pituitary produces seven hormones. These are the growth hormone 
(GH), thyroid-stimulating hormone (TSH), adrenocorticotropic hormone 
(ACTH), follicle-stimulating hormone (FSH), luteinizing hormone (LH), beta 
endorphin, and prolactin. Of the hormones of the anterior pituitary, TSH, 
ACTH, FSH, and LH are collectively referred to as tropic hormones (trope- = 
“turning”) because they turn on or off the function of other endocrine glands. 


Growth Hormone 


The endocrine system regulates the growth of the human body, protein 
synthesis, and cellular replication. A major hormone involved in this process is 
growth hormone (GH), also called somatotropin—a protein hormone 
produced and secreted by the anterior pituitary gland. Its primary function is 
anabolic; it promotes protein synthesis and tissue building through direct and 
indirect mechanisms ([link]). GH levels are controlled by the release of GHRH 
and GHIH (also known as somatostatin) from the hypothalamus. 
Hormonal Regulation of Growth 
1) Release of growth hormone: GHRH release 3) Inhibition of growth hormone: GHIH release 
¢ Hypothalamus releases * High IGF-1 levels perceived by 
growth hormone-releasing hypothalamus 
hormone (GHRH) ra fy * Growth hormone-—inhibiting 
¢ GHRH stimulates the anterior we YY hormone (GHIH) is released to 
pituitary to release growth YY inhibit GH release 


hormone (GH) * GHIH inhibits GH release in the 
anterior pituitary 


GH release Sa GH }Gase 


2a) Glucose-sparing effect: 2b) Growth effects: 2c) Diabetogenic effect: 
¢ Stimulates adipose cells ¢ Increases uptake of amino * GH stimulates liver to 
to break down stored fat, acids from the blood break down glycogen 
fueling growth effects ¢ Enhances cellular proliferation into glucose, fueling 
and reduces apoptosis growth effects 


Targets: 


Liver releases 
IGF-1, further 
Adipose cells stimulating 

growth effects 


a oe” Bone cells 


Muscle cells 
IGF-1 release 


, Nervous system 
’ cells 


Immune system 
cells 


Growth hormone (GH) directly accelerates the rate of protein 
synthesis in skeletal muscle and bones. Insulin-like growth 
factor 1 (IGF-1) is activated by growth hormone and indirectly 
supports the formation of new proteins in muscle cells and 
bone. 


A glucose-sparing effect occurs when GH stimulates lipolysis, or the 
breakdown of adipose tissue, releasing fatty acids into the blood. As a result, 
many tissues switch from glucose to fatty acids as their main energy source, 
which means that less glucose is taken up from the bloodstream. 


GH also initiates the diabetogenic effect in which GH stimulates the liver to 
break down glycogen to glucose, which is then deposited into the blood. The 
name “diabetogenic” is derived from the similarity in elevated blood glucose 
levels observed between individuals with untreated diabetes mellitus and 
individuals experiencing GH excess. Blood glucose levels rise as the result of a 
combination of glucose-sparing and diabetogenic effects. 


GH indirectly mediates growth and protein synthesis by triggering the liver and 
other tissues to produce a group of proteins called insulin-like growth factors 
(IGFs). These proteins enhance cellular proliferation and inhibit apoptosis, or 
programmed cell death. IGFs stimulate cells to increase their uptake of amino 
acids from the blood for protein synthesis. Skeletal muscle and cartilage cells 
are particularly sensitive to stimulation from IGFs. 


Dysfunction of the endocrine system’s control of growth can result in several 
disorders. For example, gigantism is a disorder in children that is caused by the 
secretion of abnormally large amounts of GH, resulting in excessive growth. A 
similar condition in adults is acromegaly, a disorder that results in the growth 
of bones in the face, hands, and feet in response to excessive levels of GH in 
individuals who have stopped growing. Abnormally low levels of GH in 
children can cause growth impairment—a disorder called pituitary dwarfism 
(also known as growth hormone deficiency). 


Thyroid-Stimulating Hormone 


The activity of the thyroid gland is regulated by thyroid-stimulating hormone 
(TSH), also called thyrotropin. TSH is released from the anterior pituitary in 
response to thyrotropin-releasing hormone (TRH) from the hypothalamus. As 
discussed shortly, it triggers the secretion of thyroid hormones by the thyroid 
gland. In a classic negative feedback loop, elevated levels of thyroid hormones 
in the bloodstream then trigger a drop in production of TRH and subsequently 
SEL. 


Adrenocorticotropic Hormone 


The adrenocorticotropic hormone (ACTH), also called corticotropin, 
stimulates the adrenal cortex (the more superficial “bark” of the adrenal glands) 
to secrete corticosteroid hormones such as cortisol. ACTH come from a 
precursor molecule known as pro-opiomelanotropin (POMC) which produces 
several biologically active molecules when cleaved, including ACTH, 
melanocyte-stimulating hormone, and the brain opioid peptides known as 
endorphins. 


The release of ACTH is regulated by the corticotropin-releasing hormone 
(CRH) from the hypothalamus in response to normal physiologic rhythms. A 
variety of stressors can also influence its release, and the role of ACTH in the 
stress response is discussed later in this chapter. 


Follicle-Stimulating Hormone and Luteinizing Hormone 


The endocrine glands secrete a variety of hormones that control the 
development and regulation of the reproductive system (these glands include 
the anterior pituitary, the adrenal cortex, and the gonads—the testes in males 
and the ovaries in females). Much of the development of the reproductive 
system occurs during puberty and is marked by the development of sex-specific 
characteristics in both male and female adolescents. Puberty is initiated by 
gonadotropin-releasing hormone (GnRH), a hormone produced and secreted by 
the hypothalamus. GnRH stimulates the anterior pituitary to secrete 
gonadotropins—hormones that regulate the function of the gonads. The levels 
of GnRH are regulated through a negative feedback loop; high levels of 
reproductive hormones inhibit the release of GnRH. Throughout life, 


gonadotropins regulate reproductive function and, in the case of women, the 
onset and cessation of reproductive capacity. 


The gonadotropins include two glycoprotein hormones: follicle-stimulating 
hormone (FSH) stimulates the production and maturation of sex cells, or 
gametes, including ova in women and sperm in men. FSH also promotes 
follicular growth; these follicles then release estrogens in the female ovaries. 
Luteinizing hormone (LH) triggers ovulation in women, as well as the 
production of estrogens and progesterone by the ovaries. LH stimulates 
production of testosterone by the male testes. 


Prolactin 


As its name implies, prolactin (PRL) promotes lactation (milk production) in 
women. During pregnancy, it contributes to development of the mammary 
glands, and after birth, it stimulates the mammary glands to produce breast 
milk. However, the effects of prolactin depend heavily upon the permissive 
effects of estrogens, progesterone, and other hormones. And as noted earlier, 
the let-down of milk occurs in response to stimulation from oxytocin. 


In a non-pregnant woman, prolactin secretion is inhibited by prolactin- 
inhibiting hormone (PIH), which is actually the neurotransmitter dopamine, and 
is released from neurons in the hypothalamus. Only during pregnancy do 
prolactin levels rise in response to prolactin-releasing hormone (PRH) from the 
hypothalamus. 


Intermediate Pituitary: Melanocyte-Stimulating Hormone 


The cells in the zone between the pituitary lobes secrete a hormone known as 
melanocyte-stimulating hormone (MSH) that is formed by cleavage of the pro- 
opiomelanocortin (POMC) precursor protein. Local production of MSH in the 
skin is responsible for melanin production in response to UV light exposure. 
The role of MSH made by the pituitary is more complicated. For instance, 
people with lighter skin generally have the same amount of MSH as people 
with darker skin. Nevertheless, this hormone is capable of darkening of the skin 
by inducing melanin production in the skin’s melanocytes. Women also show 
increased MSH production during pregnancy; in combination with estrogens, it 


can lead to darker skin pigmentation, especially the skin of the areolas and 
labia minora. [link] is a summary of the pituitary hormones and their principal 
effects. 

Major Pituitary Hormones 


Posterior Pituitary Hormones 


Releasing hormone Pituitary 
(hypothalamus) hormone Target Effects 
ADH Stores ————» Kidneys, ——_ Water balance 
ADH sweat glands, 
circulatory 
system 
- OT ——+» Female ——~ Triggers uterine 
reproductive contractions during 
system childbirth 


Anterior Pituitary Hormones 


Releasing hormone Pituitary 
(hypothalamus) hormone Target Effects 
GnRH = ——e» LH ——+* Reproductive ———® Stimulates production 
system of sex hormones by 
gonads 


GnRH ——» FSH ——® Ffeproductive ———» Stimulates production 
system of sperm and eggs 


TRH —> TSH — Thyroid gland ———* Stimulates the release 
of thyroid hormone 
(TH). TH regulates 


metabolism. 
PRH —> +~=PRL — Mammary ——_ Promotes milk 
(inhibited glands production 
by PIH) 
GHRH mY GH — Liver,bone, ——— Induces targets to 
(inhibited muscles produce insulin-like 
by GHIH) growth factors (IGF). 
IGFs stimulate body 


growth and a higher 
metabolic rate. 


CRH ——» ACTH ——+* Adrenal ——> Induces targets to 
glands produce glucocorticoids, 
which regulate 
metabolism and the 
stress response 


Major pituitary hormones and their target organs. 


Note: 


Sch 


Visit this link to watch an animation showing the role of the hypothalamus and 
the pituitary gland. Which hormone is released by the pituitary to stimulate the 
thyroid gland? 


Chapter Review 


The hypothalamus—pituitary complex is located in the diencephalon of the 
brain. The hypothalamus and the pituitary gland are connected by a structure 
called the infundibulum, which contains vasculature and nerve axons. The 
pituitary gland is divided into two distinct structures with different embryonic 
origins. The posterior lobe houses the axon terminals of hypothalamic neurons. 
It stores and releases into the bloodstream two hypothalamic hormones: 
oxytocin and antidiuretic hormone (ADH). The anterior lobe is connected to 
the hypothalamus by vasculature in the infundibulum and produces and 
secretes six hormones. Their secretion is regulated, however, by releasing and 
inhibiting hormones from the hypothalamus. The six anterior pituitary 
hormones are: growth hormone (GH), thyroid-stimulating hormone (TSH), 
adrenocorticotropic hormone (ACTH), follicle-stimulating hormone (FSH), 
luteinizing hormone (LH), and prolactin (PRL). 


Interactive Link Questions 


Exercise: 


Problem: 


Visit this link to watch an animation showing the role of the hypothalamus 
and the pituitary gland. Which hormone is released by the pituitary to 
stimulate the thyroid gland? 


Solution: 


Thyroid-stimulating hormone. 


Review Questions 
Exercise: 


Problem: 


The hypothalamus is functionally and anatomically connected to the 
posterior pituitary lobe by a bridge of 


a. blood vessels 
b. nerve axons 
c. cartilage 

d. bone 


Solution: 


B 


Exercise: 


Problem: Which of the following is an anterior pituitary hormone? 


a. ADH 

b. oxytocin 
c. TSH 

d. cortisol 


Solution: 
C 
Exercise: 
Problem:How many hormones are produced by the posterior pituitary? 


a. O 


ans 
DMN 


Solution: 


A 
Exercise: 


Problem: 


Which of the following hormones contributes to the regulation of the 
body’s fluid and electrolyte balance? 


a. adrenocorticotropic hormone 
b. antidiuretic hormone 

c. luteinizing hormone 

d. all of the above 


Solution: 


B 


Critical Thinking Questions 


Exercise: 
Problem: 


Compare and contrast the anatomical relationship of the anterior and 
posterior lobes of the pituitary gland to the hypothalamus. 


Solution: 


The anterior lobe of the pituitary gland is connected to the hypothalamus 
by vasculature, which allows regulating hormones from the hypothalamus 
to travel to the anterior pituitary. In contrast, the posterior lobe is 
connected to the hypothalamus by a bridge of nerve axons called the 


hypothalamic—hypophyseal tract, along which the hypothalamus sends 
hormones produced by hypothalamic nerve cell bodies to the posterior 
pituitary for storage and release into the circulation. 


Exercise: 


Problem: Name the target tissues for prolactin. 


Solution: 


The mammary glands are the target tissues for prolactin. 


Glossary 


acromegaly 
disorder in adults caused when abnormally high levels of GH trigger 
growth of bones in the face, hands, and feet 


adrenocorticotropic hormone (ACTH) 
anterior pituitary hormone that stimulates the adrenal cortex to secrete 
corticosteroid hormones (also called corticotropin) 


antidiuretic hormone (ADH) 
hypothalamic hormone that is stored by the posterior pituitary and that 
signals the kidneys to reabsorb water 


follicle-stimulating hormone (FSH) 
anterior pituitary hormone that stimulates the production and maturation 
of sex cells 


gigantism 
disorder in children caused when abnormally high levels of GH prompt 
excessive growth 


gonadotropins 
hormones that regulate the function of the gonads 


growth hormone (GH) 


anterior pituitary hormone that promotes tissue building and influences 
nutrient metabolism (also called somatotropin) 


hypophyseal portal system 
network of blood vessels that enables hypothalamic hormones to travel 
into the anterior lobe of the pituitary without entering the systemic 
circulation 


hypothalamus 
region of the diencephalon inferior to the thalamus that functions in neural 
and endocrine signaling 


infundibulum 
stalk containing vasculature and neural tissue that connects the pituitary 
gland to the hypothalamus (also called the pituitary stalk) 


insulin-like growth factors (IGF) 
protein that enhances cellular proliferation, inhibits apoptosis, and 
stimulates the cellular uptake of amino acids for protein synthesis 


luteinizing hormone (LH) 
anterior pituitary hormone that triggers ovulation and the production of 
ovarian hormones in females, and the production of testosterone in males 


osmoreceptor 
hypothalamic sensory receptor that is stimulated by changes in solute 
concentration (osmotic pressure) in the blood 


oxytocin 
hypothalamic hormone stored in the posterior pituitary gland and 
important in stimulating uterine contractions in labor, milk ejection during 
breastfeeding, and feelings of attachment (also produced in males) 


pituitary dwarfism 
disorder in children caused when abnormally low levels of GH result in 
growth retardation 


pituitary gland 
bean-sized organ suspended from the hypothalamus that produces, stores, 
and secretes hormones in response to hypothalamic stimulation (also 


called hypophysis) 


prolactin (PRL) 
anterior pituitary hormone that promotes development of the mammary 
glands and the production of breast milk 


thyroid-stimulating hormone (TSH) 
anterior pituitary hormone that triggers secretion of thyroid hormones by 
the thyroid gland (also called thyrotropin) 


The Thyroid Gland 
By the end of this section, you will be able to: 


¢ Describe the location and anatomy of the thyroid gland 

e Discuss the synthesis of triiodothyronine and thyroxine 

e Explain the role of thyroid hormones in the regulation of basal 
metabolism 

e Identify the hormone produced by the parafollicular cells of the 
thyroid 


A butterfly-shaped organ, the thyroid gland is located anterior to the 
trachea, just inferior to the larynx ([link]). The medial region, called the 
isthmus, is flanked by wing-shaped left and right lobes. Each of the thyroid 
lobes are embedded with parathyroid glands, primarily on their posterior 
surfaces. The tissue of the thyroid gland is composed mostly of thyroid 
follicles. The follicles are made up of a central cavity filled with a sticky 
fluid called colloid. Surrounded by a wall of epithelial follicle cells, the 
colloid is the center of thyroid hormone production, and that production is 
dependent on the hormones’ essential and unique component: iodine. 
Thyroid Gland 


Hyoid bone 


Thyroid cartilage 


Superior thyroid 
artery 

Isthmus of the 
thyroid 


Common carotid 


arteries Trachea 


Hyoid bone 
Thyroid cartilage 


Cricoid cartilage 


Right parathyroid 


Left parathyroid glands 


glands 


Left inferior thyroid 
artery 


From left subclavian _, * From right subclavian 
artery artery 
b) Posterior view 


Right inferior thyroid 
artery 


Parafollicular cell 


Colloid-containing 
follicle 


Follicle cells 
(cuboidal epithelium) 


c) Thyroid follicle cells 


The thyroid gland is located in the neck 
where it wraps around the trachea. (a) 
Anterior view of the thyroid gland. (b) 
Posterior view of the thyroid gland. (c) 


The glandular tissue is composed 
primarily of thyroid follicles. The larger 
parafollicular cells often appear within the 
matrix of follicle cells. LM x 1332. 
(Micrograph provided by the Regents of 
University of Michigan Medical School © 
2012) 


Synthesis and Release of Thyroid Hormones 


Hormones are produced in the colloid when atoms of the mineral iodine 
attach to a glycoprotein, called thyroglobulin, that is secreted into the 
colloid by the follicle cells. The following steps outline the hormones’ 
assembly: 


1. Binding of TSH to its receptors in the follicle cells of the thyroid gland 


causes the cells to actively transport iodide ions (I-) across their cell 
membrane, from the bloodstream into the cytosol. As a result, the 
concentration of iodide ions “trapped” in the follicular cells is many 
times higher than the concentration in the bloodstream. 


2. Iodide ions then move to the lumen of the follicle cells that border the 


colloid. There, the ions undergo oxidation (their negatively charged 


electrons are removed). The oxidation of two iodide ions (2 I") results 
in iodine (Ip), which passes through the follicle cell membrane into the 


colloid. 
3. In the colloid, peroxidase enzymes link the iodine to the tyrosine 


amino acids in thyroglobulin to produce two intermediaries: a tyrosine 


attached to one iodine and a tyrosine attached to two iodines. When 
one of each of these intermediaries is linked by covalent bonds, the 


resulting compound is triiodothyronine (T3), a thyroid hormone with 


three iodines. Much more commonly, two copies of the second 
intermediary bond, forming tetraiodothyronine, also known as 
thyroxine (T,), a thyroid hormone with four iodines. 


These hormones remain in the colloid center of the thyroid follicles until 
TSH stimulates endocytosis of colloid back into the follicle cells. There, 
lysosomal enzymes break apart the thyroglobulin colloid, releasing free T3 
and Ty, which diffuse across the follicle cell membrane and enter the 
bloodstream. 


In the bloodstream, less than one percent of the circulating T3 and T, 
remains unbound. This free T3 and T, can cross the lipid bilayer of cell 
membranes and be taken up by cells. The remaining 99 percent of 
circulating T3 and Ty is bound to specialized transport proteins called 
thyroxine-binding globulins (TBGs), to albumin, or to other plasma 
proteins. This “packaging” prevents their free diffusion into body cells. 
When blood levels of T3 and Ty begin to decline, bound T3 and Ty are 
released from these plasma proteins and readily cross the membrane of 
target cells. T3 is more potent than Ty, and many cells convert T, to T3 
through the removal of an iodine atom. 


Regulation of TH Synthesis 


The release of T3 and T, from the thyroid gland is regulated by thyroid- 
stimulating hormone (TSH). As shown in [link], low blood levels of T3 and 
T, stimulate the release of thyrotropin-releasing hormone (TRH) from the 
hypothalamus, which triggers secretion of TSH from the anterior pituitary. 
In turn, TSH stimulates the thyroid gland to secrete T3 and Ty. The levels of 
TRH, TSH, T3, and Ty are regulated by a negative feedback system in 
which increasing levels of T3 and T, decrease the production and secretion 
or TSH, 

Classic Negative Feedback Loop 


1) Metabolic rate and/or T; and T,4 
concentration in blood... 
Low? 


* Hypothalamus releases TRH. : 
This triggers TSH release by High? 


the pituitary. * Hypothalamus stops TRH 4) Negative feedback: 


<—— 


release ¢ Elevated T3 and T, levels 
inhibit release of TRH and TSH 


—— 


¢ Anterior pituitary stops 
TSH release 


Thyroid follicle 


T, release 
T, release 
3) Effects of T, and T, release: 
¢ Increased basal metabolic 
2) Effects of TSH release: rate of body cells 
* Triggers release of T3 and T4 * Rise in body temperature 
by thyroid follicle cells (calorigenic effect) 


A classic negative feedback loop controls the regulation of 
thyroid hormone levels. 


Functions of Thyroid Hormones 


The thyroid hormones, T3 and Ty, are often referred to as metabolic 
hormones because their levels influence the body’s basal metabolic rate, the 
amount of energy used by the body at rest. When T3 and Ty bind to 
intracellular receptors located on the mitochondria, they cause an increase 
in nutrient breakdown and the use of oxygen to produce ATP. In addition, 
T3 and T, initiate the transcription of genes involved in glucose oxidation. 


Although these mechanisms prompt cells to produce more ATP, the process 
is inefficient, and an abnormally increased level of heat is released as a 
byproduct of these reactions. This so-called calorigenic effect (calor- = 
“heat”) raises body temperature. 


Adequate levels of thyroid hormones are also required for protein synthesis 
and for fetal and childhood tissue development and growth. They are 
especially critical for normal development of the nervous system both in 
utero and in early childhood, and they continue to support neurological 
function in adults. As noted earlier, these thyroid hormones have a complex 
interrelationship with reproductive hormones, and deficiencies can 
influence libido, fertility, and other aspects of reproductive function. 
Finally, thyroid hormones increase the body’s sensitivity to catecholamines 
(epinephrine and norepinephrine) from the adrenal medulla by upregulation 
of receptors in the blood vessels. When levels of T3 and T, hormones are 
excessive, this effect accelerates the heart rate, strengthens the heartbeat, 
and increases blood pressure. Because thyroid hormones regulate 
metabolism, heat production, protein synthesis, and many other body 
functions, thyroid disorders can have severe and widespread consequences. 


Note: 

Disorders of the... 

Endocrine System: Iodine Deficiency, Hypothyroidism, and 
Hyperthyroidism 

As discussed above, dietary iodine is required for the synthesis of T3 and 
Ty. But for much of the world’s population, foods do not provide adequate 
levels of this mineral, because the amount varies according to the level in 
the soil in which the food was grown, as well as the irrigation and 
fertilizers used. Marine fish and shrimp tend to have high levels because 
they concentrate iodine from seawater, but many people in landlocked 
regions lack access to seafood. Thus, the primary source of dietary iodine 
in many countries is iodized salt. Fortification of salt with iodine began in 
the United States in 1924, and international efforts to iodize salt in the 
world’s poorest nations continue today. 


Dietary iodine deficiency can result in the impaired ability to synthesize T3 
and Ty, leading to a variety of severe disorders. When T3 and T,4 cannot be 
produced, TSH is secreted in increasing amounts. As a result of this 
hyperstimulation, thyroglobulin accumulates in the thyroid gland follicles, 
increasing their deposits of colloid. The accumulation of colloid increases 
the overall size of the thyroid gland, a condition called a goiter ([link]). A 
goiter is only a visible indication of the deficiency. Other iodine deficiency 
disorders include impaired growth and development, decreased fertility, 
and prenatal and infant death. Moreover, iodine deficiency is the primary 
cause of preventable mental retardation worldwide. Neonatal 
hypothyroidism (cretinism) is characterized by cognitive deficits, short 
stature, and sometimes deafness and muteness in children and adults born 
to mothers who were iodine-deficient during pregnancy. 

Goiter 


(credit: “Almazi”/Wikimedia Commons) 


In areas of the world with access to iodized salt, dietary deficiency is rare. 
Instead, inflammation of the thyroid gland is the more common cause of 
low blood levels of thyroid hormones. Called hypothyroidism, the 
condition is characterized by a low metabolic rate, weight gain, cold 
extremities, constipation, reduced libido, menstrual irregularities, and 
reduced mental activity. In contrast, hyperthyroidism—an abnormally 


elevated blood level of thyroid hormones—is often caused by a pituitary or 
thyroid tumor. In Graves’ disease, the hyperthyroid state results from an 
autoimmune reaction in which antibodies overstimulate the follicle cells of 
the thyroid gland. Hyperthyroidism can lead to an increased metabolic rate, 
excessive body heat and sweating, diarrhea, weight loss, tremors, and 
increased heart rate. The person’s eyes may bulge (called exophthalmos) as 
antibodies produce inflammation in the soft tissues of the orbits. The 
person may also develop a goiter. 


Calcitonin 


The thyroid gland also secretes a hormone called calcitonin that is 
produced by the parafollicular cells (also called C cells) that stud the tissue 
between distinct follicles. Calcitonin is released in response to a rise in 
blood calcium levels. It appears to have a function in decreasing blood 
calcium concentrations by: 


e Inhibiting the activity of osteoclasts, bone cells that release calcium 
into the circulation by degrading bone matrix 

e Increasing osteoblastic activity 

e Decreasing calcium absorption in the intestines 

e Increasing calcium loss in the urine 


However, these functions are usually not significant in maintaining calcium 
homeostasis, so the importance of calcitonin is not entirely understood. 
Pharmaceutical preparations of calcitonin are sometimes prescribed to 
reduce osteoclast activity in people with osteoporosis and to reduce the 
degradation of cartilage in people with osteoarthritis. The hormones 
secreted by thyroid are summarized in [link]. 


Thyroid Hormones 


Chemical 
Associated hormones class Effect 
Thyroxine (Ta), melee Stimulate basal 
triiodothyronine (T3) metabolic rate 
2+ 
Calcitonin Peptide Reduces blood Ca 
levels 


Of course, calcium is critical for many other biological processes. It is a 
second messenger in many signaling pathways, and is essential for muscle 
contraction, nerve impulse transmission, and blood clotting. Given these 
roles, it is not surprising that blood calcium levels are tightly regulated by 
the endocrine system. The organs involved in the regulation are the 
parathyroid glands. 


Chapter Review 


The thyroid gland is a butterfly-shaped organ located in the neck anterior to 
the trachea. Its hormones regulate basal metabolism, oxygen use, nutrient 
metabolism, the production of ATP, and calcium homeostasis. They also 
contribute to protein synthesis and the normal growth and development of 
body tissues, including maturation of the nervous system, and they increase 
the body’s sensitivity to catecholamines. The thyroid hormones 
triiodothyronine (T3) and thyroxine (T4) are produced and secreted by the 
thyroid gland in response to thyroid-stimulating hormone (TSH) from the 
anterior pituitary. Synthesis of the amino acid—derived T3 and T, hormones 
requires iodine. Insufficient amounts of iodine in the diet can lead to goiter, 
cretinism, and many other disorders. 


Review Questions 


Exercise: 


Problem: 
Which of the following statements about the thyroid gland is true? 


a. It is located anterior to the trachea and inferior to the larynx. 
b. The parathyroid glands are embedded within it. 

c. It manufactures three hormones. 

d. all of the above 


Solution: 


D 
Exercise: 


Problem: 
The secretion of thyroid hormones is controlled by 


a. TSH from the hypothalamus 

b. TSH from the anterior pituitary 

c. thyroxine from the anterior pituitary 

d. thyroglobulin from the thyroid’s parafollicular cells 


Solution: 


B 


Exercise: 


Problem:The development of a goiter indicates that 


a. the anterior pituitary is abnormally enlarged 

b. there is hypertrophy of the thyroid’s follicle cells 

c. there is an excessive accumulation of colloid in the thyroid 
follicles 

d. the anterior pituitary is secreting excessive growth hormone 


Solution: 


C 
Exercise: 


Problem: 


Iodide ions cross from the bloodstream into follicle cells via 


a. simple diffusion 

b. facilitated diffusion 
c. active transport 

d. osmosis 


Solution: 


C 


Critical Thinking Questions 


Exercise: 
Problem: 


Explain why maternal iodine deficiency might lead to neurological 
impairment in the fetus. 


Solution: 


Iodine deficiency in a pregnant woman would also deprive the fetus. 
Iodine is required for the synthesis of thyroid hormones, which 
contribute to fetal growth and development, including maturation of 
the nervous system. Insufficient amounts would impair these functions. 


Exercise: 


Problem: 


Define hyperthyroidism and explain why one of its symptoms is 
weight loss. 


Solution: 


Hyperthyroidism is an abnormally elevated blood level of thyroid 
hormones due to an overproduction of T3 and T,. An individual with 
hyperthyroidism is likely to lose weight because one of the primary 
roles of thyroid hormones is to increase the body’s basal metabolic 
rate, increasing the breakdown of nutrients and the production of ATP. 


Glossary 


calcitonin 
peptide hormone produced and secreted by the parafollicular cells (C 
cells) of the thyroid gland that functions to decrease blood calcium 
levels 


colloid 
viscous fluid in the central cavity of thyroid follicles, containing the 
glycoprotein thyroglobulin 


goiter 
enlargement of the thyroid gland either as a result of iodine deficiency 
or hyperthyroidism 


hyperthyroidism 
clinically abnormal, elevated level of thyroid hormone in the blood; 
characterized by an increased metabolic rate, excess body heat, 
sweating, diarrhea, weight loss, and increased heart rate 


hypothyroidism 
clinically abnormal, low level of thyroid hormone in the blood; 
characterized by low metabolic rate, weight gain, cold extremities, 
constipation, and reduced mental activity 


neonatal hypothyroidism 
condition characterized by cognitive deficits, short stature, and other 
signs and symptoms in people born to women who were iodine- 
deficient during pregnancy 


thyroid gland 
large endocrine gland responsible for the synthesis of thyroid 
hormones 


thyroxine 
(also, tetraiodothyronine, T4) amino acid—derived thyroid hormone that 
is more abundant but less potent than T3 and often converted to T3 by 
target cells 


triiodothyronine 
(also, T3) amino acid—derived thyroid hormone that is less abundant 
but more potent than T, 


The Parathyroid Glands 
By the end of this section, you will be able to: 


¢ Describe the location and structure of the parathyroid glands 
e Describe the hormonal control of blood calcium levels 
e Discuss the physiological response of parathyroid dysfunction 


The parathyroid glands are tiny, round structures usually found embedded 
in the posterior surface of the thyroid gland ([link]). A thick connective 
tissue capsule separates the glands from the thyroid tissue. Most people 
have four parathyroid glands, but occasionally there are more in tissues of 
the neck or chest. The function of one type of parathyroid cells, the oxyphil 
cells, is not clear. The primary functional cells of the parathyroid glands are 
the chief cells. These epithelial cells produce and secrete the parathyroid 
hormone (PTH), the major hormone involved in the regulation of blood 
calcium levels. 
Parathyroid Glands 


pa 
Hyoid bone oa Ly 
Thyroid y| 4 


cartilage 


Oxyphil cells 


Cricoid 


i Blood vessel 
cartilage 


- Wit P 7 ; y: OT ed. Oe = 1 Parathyroid 
parathyroid 


parathyroid 
glands 


glands 


a) Thyroid gland, posterior view b) Micrograph of parathyroid tissue 


The small parathyroid glands are embedded in the posterior 
surface of the thyroid gland. LM x 760. (Micrograph provided 
by the Regents of University of Michigan Medical School © 
2012) 


Note: 


mss" OPENStax COLLEGE 
F : ca 


[aS 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. 


The parathyroid glands produce and secrete PTH, a peptide hormone, in 
response to low blood calcium levels ({link]). PTH secretion causes the 
release of calcium from the bones by stimulating osteoclasts, which secrete 
enzymes that degrade bone and release calcium into the interstitial fluid. 
PTH also inhibits osteoblasts, the cells involved in bone deposition, thereby 
sparing blood calcium. PTH causes increased reabsorption of calcium (and 
magnesium) in the kidney tubules from the urine filtrate. In addition, PTH 
initiates the production of the steroid hormone calcitriol (also known as 
1,25-dihydroxyvitamin D), which is the active form of vitamin Ds, in the 
kidneys. Calcitriol then stimulates increased absorption of dietary calcium 
by the intestines. A negative feedback loop regulates the levels of PTH, 
with rising blood calcium levels inhibiting further release of PTH. 
Parathyroid Hormone in Maintaining Blood Calcium Homeostasis 


\ 2s Osteoclasts 

Ss : 
1) Blood calcium Compact bone 
concentration drops 


6) Effects of calcitonin on bone: 
+ Stimulates osteoblasts 
+ Inhibits osteoclasts 


+ Calcium is removed from blood and used to build bone 


2) Release of PTH: 5) Calcitonin release: 
+ Chief cells of the parathyroid gland release parathyroid 
hormone (PTH). 


+ High concentrations of calcium stimulate parafollicular cells 
in the thyroid to release calcitonin. 


i 4 C 
¥ rr 
4 2 TA EAT vy 


om 


4) Blood calcium levels increase 


3a) Effects of PTH on bone: 3b) Effects of PTH on kidneys: 3c) Effects of calcitriol on intestine: 
+ Inhibits osteoblasts + PTH stimulates kidney tubule cells to + Stimulates intestines to absorb 
+ Stimulates osteoclasts recover waste calcium from the urine. calcium from digesting food 
+ Bone is broken down, releasing + PTH stimulates kidney tubule cells 
calcium ions into bloodstream to release calcitriol. 


i at Intestinal lumen 
Kidney _ Interstitial 
Urine tubule cells fluid 


+ Food 


— . Intestinal cells 
Compact / a A 
bone = 


| Intestinal 
IF - : connective tissue 
Osteoclasts = | with blood supply 


PT 


Osteoblasts 


Parathyroid hormone increases blood calcium levels when they 
drop too low. Conversely, calcitonin, which is released from 
the thyroid gland, decreases blood calcium levels when they 

become too high. These two mechanisms constantly maintain 

blood calcium concentration at homeostasis. 


Abnormally high activity of the parathyroid gland can cause 
hyperparathyroidism, a disorder caused by an overproduction of PTH that 
results in excessive calcium reabsorption from bone. Hyperparathyroidism 
can significantly decrease bone density, leading to spontaneous fractures or 
deformities. As blood calcium levels rise, cell membrane permeability to 
sodium is decreased, and the responsiveness of the nervous system is 
reduced. At the same time, calcium deposits may collect in the body’s 
tissues and organs, impairing their functioning. 


In contrast, abnormally low blood calcium levels may be caused by 
parathyroid hormone deficiency, called hypoparathyroidism, which may 
develop following injury or surgery involving the thyroid gland. Low blood 
calcium increases membrane permeability to sodium, resulting in muscle 
twitching, cramping, spasms, or convulsions. Severe deficits can paralyze 
muscles, including those involved in breathing, and can be fatal. 


When blood calcium levels are high, calcitonin is produced and secreted by 
the parafollicular cells of the thyroid gland. As discussed earlier, calcitonin 
inhibits the activity of osteoclasts, reduces the absorption of dietary calcium 
in the intestine, and signals the kidneys to reabsorb less calcium, resulting 
in larger amounts of calcium excreted in the urine. 


Chapter Review 


Calcium is required for a variety of important physiologic processes, 
including neuromuscular functioning; thus, blood calcium levels are closely 
regulated. The parathyroid glands are small structures located on the 
posterior thyroid gland that produce parathyroid hormone (PTH), which 
regulates blood calcium levels. Low blood calcium levels cause the 
production and secretion of PTH. In contrast, elevated blood calcium levels 
inhibit secretion of PTH and trigger secretion of the thyroid hormone 
calcitonin. Underproduction of PTH can result in hypoparathyroidism. In 
contrast, overproduction of PTH can result in hyperparathyroidism. 


Review Questions 


Exercise: 


Problem: 
When blood calcium levels are low, PTH stimulates 


a. urinary excretion of calcium by the kidneys 

b. a reduction in calcium absorption from the intestines 
c. the activity of osteoblasts 

d. the activity of osteoclasts 


Solution: 


D 
Exercise: 


Problem: 
Which of the following can result from hyperparathyroidism? 


a. increased bone deposition 
b. fractures 

c. convulsions 

d. all of the above 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


Describe the role of negative feedback in the function of the 
parathyroid gland. 


Solution: 


The production and secretion of PTH is regulated by a negative 
feedback loop. Low blood calcium levels initiate the production and 
secretion of PTH. PTH increases bone resorption, calcium absorption 
from the intestines, and calcium reabsorption by the kidneys. As a 
result, blood calcium levels begin to rise. This, in turn, inhibits the 
further production and secretion of PTH. 


Exercise: 
Problem: 


Explain why someone with a parathyroid gland tumor might develop 
kidney stones. 


Solution: 


A parathyroid gland tumor can prompt hypersecretion of PTH. This 
can raise blood calcium levels so excessively that calcium deposits 
begin to accumulate throughout the body, including in the kidney 
tubules, where they are referred to as kidney stones. 


Glossary 


hyperparathyroidism 
disorder caused by overproduction of PTH that results in abnormally 
elevated blood calcium 


hypoparathyroidism 
disorder caused by underproduction of PTH that results in abnormally 
low blood calcium 


parathyroid glands 
small, round glands embedded in the posterior thyroid gland that 
produce parathyroid hormone (PTH) 


parathyroid hormone (PTH) 


peptide hormone produced and secreted by the parathyroid glands in 
response to low blood calcium levels 


The Adrenal Glands 
By the end of this section, you will be able to: 


¢ Describe the location and structure of the adrenal glands 
e Identify the hormones produced by the adrenal cortex and adrenal 
medulla, and summarize their target cells and effects 


The adrenal glands are wedges of glandular and neuroendocrine tissue 
adhering to the top of the kidneys by a fibrous capsule ({link]). The adrenal 
glands have a rich blood supply and experience one of the highest rates of 
blood flow in the body. They are served by several arteries branching off 
the aorta, including the suprarenal and renal arteries. Blood flows to each 
adrenal gland at the adrenal cortex and then drains into the adrenal medulla. 
Adrenal hormones are released into the circulation via the left and right 
suprarenal veins. 

Adrenal Glands 


Connective tissue | 
capsule 


Tissue area Hormones released Examples 


‘+ Zona glomerulosa ——*> Mineralcorticoids 


eee f eh ‘ Se hs cortex) (regulate mineral 
4 
pes - oe 4} 


Aldosterone 
balance) 

Zona fasciculata ———® Glucocorticoids Cortisol 

(adrenal cortex) (regulate glucose Corticosterone 


metabolism) Cortisone 


Zona reticularis ———- Androgens A 
a Dehydroepian- 
(adrenal cortex) (stimulate rap sabe 
masculinization) 


Adrenal medulla ——— Stress hormones Epinephrine 
(stimulate Norepinephrine 
sympathetic ANS) 


Adrenal gland 


Superior surface of 
kidney 


Both adrenal glands sit atop the kidneys and are composed of 
an outer cortex and an inner medulla, all surrounded by a 
connective tissue capsule. The cortex can be subdivided into 
additional zones, all of which produce different types of 
hormones. LM x 204. (Micrograph provided by the Regents of 
University of Michigan Medical School © 2012) 


Note: 


od 


es 


—_s 
mss’ OPENStax COLLEGE 
— , 7 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. 


The adrenal gland consists of an outer cortex of glandular tissue and an 
inner medulla of nervous tissue. The cortex itself is divided into three 
zones: the zona glomerulosa, the zona fasciculata, and the zona 
reticularis. Each region secretes its own set of hormones. 


The adrenal cortex, as a component of the hypothalamic-pituitary-adrenal 
(HPA) axis, secretes steroid hormones important for the regulation of the 
long-term stress response, blood pressure and blood volume, nutrient uptake 
and storage, fluid and electrolyte balance, and inflammation. The HPA axis 
involves the stimulation of hormone release of adrenocorticotropic hormone 
(ACTH) from the pituitary by the hypothalamus. ACTH then stimulates the 
adrenal cortex to produce the hormone cortisol. This pathway will be 
discussed in more detail below. 


The adrenal medulla is neuroendocrine tissue composed of postganglionic 
sympathetic nervous system (SNS) neurons. It is really an extension of the 
autonomic nervous system, which regulates homeostasis in the body. The 
sympathomedullary (SAM) pathway involves the stimulation of the 
medulla by impulses from the hypothalamus via neurons from the thoracic 
spinal cord. The medulla is stimulated to secrete the amine hormones 
epinephrine and norepinephrine. 


One of the major functions of the adrenal gland is to respond to stress. 
Stress can be either physical or psychological or both. Physical stresses 
include exposing the body to injury, walking outside in cold and wet 
conditions without a coat on, or malnutrition. Psychological stresses include 


the perception of a physical threat, a fight with a loved one, or just a bad 
day at school. 


The body responds in different ways to short-term stress and long-term 
stress following a pattern known as the general adaptation syndrome 
(GAS). Stage one of GAS is called the alarm reaction. This is short-term 
stress, the fight-or-flight response, mediated by the hormones epinephrine 
and norepinephrine from the adrenal medulla via the SAM pathway. Their 
function is to prepare the body for extreme physical exertion. Once this 
stress is relieved, the body quickly returns to normal. The section on the 
adrenal medulla covers this response in more detail. 


If the stress is not soon relieved, the body adapts to the stress in the second 
stage called the stage of resistance. If a person is starving for example, the 
body may send signals to the gastrointestinal tract to maximize the 
absorption of nutrients from food. 


If the stress continues for a longer term however, the body responds with 
symptoms quite different than the fight-or-flight response. During the stage 
of exhaustion, individuals may begin to suffer depression, the suppression 
of their immune response, severe fatigue, or even a fatal heart attack. These 
symptoms are mediated by the hormones of the adrenal cortex, especially 
cortisol, released as a result of signals from the HPA axis. 


Adrenal hormones also have several non-stress-related functions, including 


the increase of blood sodium and glucose levels, which will be described in 
detail below. 


Adrenal Cortex 
The adrenal cortex consists of multiple layers of lipid-storing cells that 


occur in three structurally distinct regions. Each of these regions produces 
different hormones. 


Note: 


. or 
mss’ OPENStAX COLLEGE 


Visit this link to view an animation describing the location and function of 
the adrenal glands. Which hormone produced by the adrenal glands is 
responsible for the mobilization of energy stores? 


Hormones of the Zona Glomerulosa 


The most superficial region of the adrenal cortex is the zona glomerulosa, 
which produces a group of hormones collectively referred to as 
mineralocorticoids because of their effect on body minerals, especially 
sodium and potassium. These hormones are essential for fluid and 
electrolyte balance. 


Aldosterone is the major mineralocorticoid. It is important in the regulation 
of the concentration of sodium and potassium ions in urine, sweat, and 
saliva. For example, it is released in response to elevated blood K™, low 
blood Na‘, low blood pressure, or low blood volume. In response, 
aldosterone increases the excretion of K” and the retention of Na‘, which in 
turn increases blood volume and blood pressure. Its secretion is prompted 
when CRH from the hypothalamus triggers ACTH release from the anterior 
pituitary. 


Aldosterone is also a key component of the renin-angiotensin-aldosterone 
system (RAAS) in which specialized cells of the kidneys secrete the 
enzyme renin in response to low blood volume or low blood pressure. 
Renin then catalyzes the conversion of the blood protein angiotensinogen, 
produced by the liver, to the hormone angiotensin I. Angiotensin I is 
converted in the lungs to angiotensin II by angiotensin-converting enzyme 
(ACE). Angiotensin II has three major functions: 


1. Initiating vasoconstriction of the arterioles, decreasing blood flow 


2. Stimulating kidney tubules to reabsorb NaCl and water, increasing 
blood volume 

3. Signaling the adrenal cortex to secrete aldosterone, the effects of 
which further contribute to fluid retention, restoring blood pressure 
and blood volume 


For individuals with hypertension, or high blood pressure, drugs are 
available that block the production of angiotensin IT. These drugs, known as 
ACE inhibitors, block the ACE enzyme from converting angiotensin I to 
angiotensin II, thus mitigating the latter’s ability to increase blood pressure. 


Hormones of the Zona Fasciculata 


The intermediate region of the adrenal cortex is the zona fasciculata, named 
as such because the cells form small fascicles (bundles) separated by tiny 
blood vessels. The cells of the zona fasciculata produce hormones called 
glucocorticoids because of their role in glucose metabolism. The most 
important of these is cortisol, some of which the liver converts to cortisone. 
A glucocorticoid produced in much smaller amounts is corticosterone. In 
response to long-term stressors, the hypothalamus secretes CRH, which in 
turn triggers the release of ACTH by the anterior pituitary. ACTH triggers 
the release of the glucocorticoids. Their overall effect is to inhibit tissue 
building while stimulating the breakdown of stored nutrients to maintain 
adequate fuel supplies. In conditions of long-term stress, for example, 
cortisol promotes the catabolism of glycogen to glucose, the catabolism of 
stored triglycerides into fatty acids and glycerol, and the catabolism of 
muscle proteins into amino acids. These raw materials can then be used to 
synthesize additional glucose and ketones for use as body fuels. The 
hippocampus, which is part of the temporal lobe of the cerebral cortices and 
important in memory formation, is highly sensitive to stress levels because 
of its many glucocorticoid receptors. 


You are probably familiar with prescription and over-the-counter 
medications containing glucocorticoids, such as cortisone injections into 
inflamed joints, prednisone tablets and steroid-based inhalers used to 
manage severe asthma, and hydrocortisone creams applied to relieve itchy 


skin rashes. These drugs reflect another role of cortisol—the 
downregulation of the immune system, which inhibits the inflammatory 
response. 


Hormones of the Zona Reticularis 


The deepest region of the adrenal cortex is the zona reticularis, which 
produces small amounts of a class of steroid sex hormones called 
androgens. During puberty and most of adulthood, androgens are produced 
in the gonads. The androgens produced in the zona reticularis supplement 
the gonadal androgens. They are produced in response to ACTH from the 
anterior pituitary and are converted in the tissues to testosterone or 
estrogens. In adult women, they may contribute to the sex drive, but their 
function in adult men is not well understood. In post-menopausal women, 
as the functions of the ovaries decline, the main source of estrogens 
becomes the androgens produced by the zona reticularis. 


Adrenal Medulla 


As noted earlier, the adrenal cortex releases glucocorticoids in response to 
long-term stress such as severe illness. In contrast, the adrenal medulla 
releases its hormones in response to acute, short-term stress mediated by the 
sympathetic nervous system (SNS). 


The medullary tissue is composed of unique postganglionic SNS neurons 
called chromaffin cells, which are large and irregularly shaped, and 
produce the neurotransmitters epinephrine (also called adrenaline) and 
norepinephrine (or noradrenaline). Epinephrine is produced in greater 
quantities—approximately a 4 to 1 ratio with norepinephrine—and is the 
more powerful hormone. Because the chromaffin cells release epinephrine 
and norepinephrine into the systemic circulation, where they travel widely 
and exert effects on distant cells, they are considered hormones. Derived 
from the amino acid tyrosine, they are chemically classified as 
catecholamines. 


The secretion of medullary epinephrine and norepinephrine is controlled by 
a neural pathway that originates from the hypothalamus in response to 
danger or stress (the SAM pathway). Both epinephrine and norepinephrine 
signal the liver and skeletal muscle cells to convert glycogen into glucose, 
resulting in increased blood glucose levels. These hormones increase the 
heart rate, pulse, and blood pressure to prepare the body to fight the 
perceived threat or flee from it. In addition, the pathway dilates the airways, 
raising blood oxygen levels. It also prompts vasodilation, further increasing 
the oxygenation of important organs such as the lungs, brain, heart, and 
skeletal muscle. At the same time, it triggers vasoconstriction to blood 
vessels serving less essential organs such as the gastrointestinal tract, 
kidneys, and skin, and downregulates some components of the immune 
system. Other effects include a dry mouth, loss of appetite, pupil dilation, 
and a loss of peripheral vision. The major hormones of the adrenal glands 
are summarized in [link]. 


Hormones of the Adrenal Glands 


Adrenal Associated Chemical 

gland hormones class Effect 

Adrenal iaeiaiane Sirota Dice’ blood 

cortex Na’ levels 

Adrenal Cortisol, ; Increase blood 
corticosterone, Steroid 

cortex ; glucose levels 
cortisone 


Stimulate fight- 
Amine or-flight 
response 


Adrenal Epinephrine, 
medulla norepinephrine 


Disorders Involving the Adrenal Glands 


Several disorders are caused by the dysregulation of the hormones produced 
by the adrenal glands. For example, Cushing’s disease is a disorder 
characterized by high blood glucose levels and the accumulation of lipid 
deposits on the face and neck. It is caused by hypersecretion of cortisol. The 
most common source of Cushing’s disease is a pituitary tumor that secretes 
cortisol or ACTH in abnormally high amounts. Other common signs of 
Cushing’s disease include the development of a moon-shaped face, a 
buffalo hump on the back of the neck, rapid weight gain, and hair loss. 
Chronically elevated glucose levels are also associated with an elevated risk 
of developing type 2 diabetes. In addition to hyperglycemia, chronically 
elevated glucocorticoids compromise immunity, resistance to infection, and 
memory, and can result in rapid weight gain and hair loss. 


In contrast, the hyposecretion of corticosteroids can result in Addison’s 
disease, a rare disorder that causes low blood glucose levels and low blood 
sodium levels. The signs and symptoms of Addison’s disease are vague and 
are typical of other disorders as well, making diagnosis difficult. They may 
include general weakness, abdominal pain, weight loss, nausea, vomiting, 
sweating, and cravings for salty food. 


Chapter Review 


The adrenal glands, located superior to each kidney, consist of two regions: 
the adrenal cortex and adrenal medulla. The adrenal cortex—the outer layer 
of the gland—produces mineralocorticoids, glucocorticoids, and androgens. 
The adrenal medulla at the core of the gland produces epinephrine and 
norepinephrine. 


The adrenal glands mediate a short-term stress response and a long-term 
stress response. A perceived threat results in the secretion of epinephrine 
and norepinephrine from the adrenal medulla, which mediate the fight-or- 
flight response. The long-term stress response is mediated by the secretion 
of CRH from the hypothalamus, which triggers ACTH, which in turn 
stimulates the secretion of corticosteroids from the adrenal cortex. The 


mineralocorticoids, chiefly aldosterone, cause sodium and fluid retention, 
which increases blood volume and blood pressure. 


Interactive Link Questions 


Exercise: 


Problem: 
Visit this link to view an animation describing the location and 
function of the adrenal glands. Which hormone produced by the 


adrenal glands is responsible for mobilization of energy stores? 


Solution: 


Cortisol. 


Review Questions 


Exercise: 


Problem:The adrenal glands are attached superiorly to which organ? 


a. thyroid 

b. liver 

c. kidneys 

d. hypothalamus 


Solution: 


C 


Exercise: 


Problem: What secretory cell type is found in the adrenal medulla? 


a. chromaffin cells 
b. neuroglial cells 
c. follicle cells 

d. oxyphil cells 


Solution: 


A 


Exercise: 


Problem: Cushing’s disease is a disorder caused by 


a. abnormally low levels of cortisol 
b. abnormally high levels of cortisol 
c. abnormally low levels of aldosterone 
d. abnormally high levels of aldosterone 


Solution: 


B 
Exercise: 


Problem: 


Which of the following responses s not part of the fight-or-flight 
response? 


a. pupil dilation 
b. increased oxygen supply to the lungs 


c. suppressed digestion 
d. reduced mental activity 


Solution: 


D 


Critical Thinking Questions 


Exercise: 
Problem: 


What are the three regions of the adrenal cortex and what hormones do 
they produce? 


Solution: 


The outer region is the zona glomerulosa, which produces 
mineralocorticoids such as aldosterone; the next region is the zona 
fasciculata, which produces glucocorticoids such as cortisol; the inner 
region is the zona reticularis, which produces androgens. 


Exercise: 
Problem: 


If innervation to the adrenal medulla were disrupted, what would be 
the physiological outcome? 


Solution: 


Damage to the innervation of the adrenal medulla would prevent the 
adrenal glands from responding to the hypothalamus during the fight- 
or-flight response. Therefore, the response would be reduced. 


Exercise: 


Problem: 
Compare and contrast the short-term and long-term stress response. 
Solution: 


The short-term stress response involves the hormones epinephrine and 
norepinephrine, which work to increase the oxygen supply to organs 


important for extreme muscular action such as the brain, lungs, and 
muscles. In the long-term stress response, the hormone cortisol is 
involved in catabolism of glycogen stores, proteins, and triglycerides, 
glucose and ketone synthesis, and downregulation of the immune 
system. 


Glossary 


adrenal cortex 
outer region of the adrenal glands consisting of multiple layers of 
epithelial cells and capillary networks that produces mineralocorticoids 
and glucocorticoids 


adrenal glands 
endocrine glands located at the top of each kidney that are important 
for the regulation of the stress response, blood pressure and blood 
volume, water homeostasis, and electrolyte levels 


adrenal medulla 
inner layer of the adrenal glands that plays an important role in the 
stress response by producing epinephrine and norepinephrine 


angiotensin-converting enzyme 
the enzyme that converts angiotensin I to angiotensin IT 


alarm reaction 
the short-term stress, or the fight-or-flight response, of stage one of the 
general adaptation syndrome mediated by the hormones epinephrine 
and norepinephrine 


aldosterone 
hormone produced and secreted by the adrenal cortex that stimulates 
sodium and fluid retention and increases blood volume and blood 
pressure 


chromaffin 
neuroendocrine cells of the adrenal medulla 


cortisol 
glucocorticoid important in gluconeogenesis, the catabolism of 
glycogen, and downregulation of the immune system 


epinephrine 
primary and most potent catecholamine hormone secreted by the 
adrenal medulla in response to short-term stress; also called adrenaline 


general adaptation syndrome (GAS) 
the human body’s three-stage response pattern to short- and long-term 
stress 


glucocorticoids 
hormones produced by the zona fasciculata of the adrenal cortex that 
influence glucose metabolism 


mineralocorticoids 
hormones produced by the zona glomerulosa cells of the adrenal 
cortex that influence fluid and electrolyte balance 


norepinephrine 
secondary catecholamine hormone secreted by the adrenal medulla in 
response to short-term stress; also called noradrenaline 


stage of exhaustion 
stage three of the general adaptation syndrome; the body’s long-term 
response to stress mediated by the hormones of the adrenal cortex 


stage of resistance 
stage two of the general adaptation syndrome; the body’s continued 
response to stress after stage one diminishes 


zona fasciculata 
intermediate region of the adrenal cortex that produce hormones called 
glucocorticoids 


zona glomerulosa 


most superficial region of the adrenal cortex, which produces the 
hormones collectively referred to as mineralocorticoids 


zona reticularis 
deepest region of the adrenal cortex, which produces the steroid sex 
hormones called androgens 


The Pineal Gland 
By the end of this section, you will be able to: 


e Describe the location and structure of the pineal gland 
e Discuss the function of melatonin 


Recall that the hypothalamus, part of the diencephalon of the brain, sits 
inferior and somewhat anterior to the thalamus. Inferior but somewhat 
posterior to the thalamus is the pineal gland, a tiny endocrine gland whose 
functions are not entirely clear. The pinealocyte cells that make up the 
pineal gland are known to produce and secrete the amine hormone 
melatonin, which is derived from serotonin. 


The secretion of melatonin varies according to the level of light received 
from the environment. When photons of light stimulate the retinas of the 
eyes, a nerve impulse is sent to a region of the hypothalamus called the 
suprachiasmatic nucleus (SCN), which is important in regulating biological 
rhythms. From the SCN, the nerve signal is carried to the spinal cord and 
eventually to the pineal gland, where the production of melatonin is 
inhibited. As a result, blood levels of melatonin fall, promoting 
wakefulness. In contrast, as light levels decline—such as during the evening 
—melatonin production increases, boosting blood levels and causing 
drowsiness. 


Note: 


openstax COLLEGE” 


: 


2 


Visit this link to view an animation describing the function of the hormone 
melatonin. What should you avoid doing in the middle of your sleep cycle 
that would lower melatonin? 


The secretion of melatonin may influence the body’s circadian rhythms, the 
dark-light fluctuations that affect not only sleepiness and wakefulness, but 
also appetite and body temperature. Interestingly, children have higher 
melatonin levels than adults, which may prevent the release of 
gonadotropins from the anterior pituitary, thereby inhibiting the onset of 
puberty. Finally, an antioxidant role of melatonin is the subject of current 
research. 


Jet lag occurs when a person travels across several time zones and feels 
sleepy during the day or wakeful at night. Traveling across multiple time 
zones significantly disturbs the light-dark cycle regulated by melatonin. It 
can take up to several days for melatonin synthesis to adjust to the light- 
dark patterns in the new environment, resulting in jet lag. Some air travelers 
take melatonin supplements to induce sleep. 


Chapter Review 


The pineal gland is an endocrine structure of the diencephalon of the brain, 
and is located inferior and posterior to the thalamus. It is made up of 
pinealocytes. These cells produce and secrete the hormone melatonin in 
response to low light levels. High blood levels of melatonin induce 
drowsiness. Jet lag, caused by traveling across several time zones, occurs 
because melatonin synthesis takes several days to readjust to the light-dark 
patterns in the new environment. 


Interactive Link Questions 


Exercise: 


Problem: 
Visit this link to view an animation describing the function of the 
hormone melatonin. What should you avoid doing in the middle of 


your sleep cycle that would lower melatonin? 


Solution: 


Turning on the lights. 


Review Questions 


Exercise: 


Problem: What cells secrete melatonin? 
a. melanocytes 
b. pinealocytes 


c. suprachiasmatic nucleus cells 
d. retinal cells 


Solution: 


B 


Exercise: 


Problem: The production of melatonin is inhibited by 
a. declining levels of light 
b. exposure to bright light 
c. the secretion of serotonin 
d. the activity of pinealocytes 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


Seasonal affective disorder (SAD) is a mood disorder characterized by, 
among other symptoms, increased appetite, sluggishness, and 
increased sleepiness. It occurs most commonly during the winter 
months, especially in regions with long winter nights. Propose a role 
for melatonin in SAD and a possible non-drug therapy. 


Solution: 


SAD is thought to occur in part because low levels and duration of 
sunlight allow excessive and prolonged secretion of melatonin. Light 
therapy—daytime exposure to very bright lighting—is one common 
therapy. 


Exercise: 
Problem: 
Retinitis pigmentosa (RP) is a disease that causes deterioration of the 


retinas of the eyes. Describe the impact RP would have on melatonin 
levels. 


Solution: 


The retina is important for melatonin production because it senses 
light. Bright light inhibits the production of melatonin, whereas low 
light levels promote the production of melatonin. Therefore, 
deterioration of the retinas would most likely disturb the sleep-wake 
pattern because melatonin production would be elevated. 


Glossary 


melatonin 
amino acid—derived hormone that is secreted in response to low light 
and causes drowsiness 


pineal gland 


endocrine gland that secretes melatonin, which is important in 
regulating the sleep-wake cycle 


pinealocyte 
cell of the pineal gland that produces and secretes the hormone 
melatonin 


Gonadal and Placental Hormones 
By the end of this section, you will be able to: 


e Identify the most important hormones produced by the testes and 
ovaries 
e Name the hormones produced by the placenta and state their functions 


This section briefly discusses the hormonal role of the gonads—the male 
testes and female ovaries—which produce the sex cells (sperm and ova) and 
secrete the gonadal hormones. The roles of the gonadotropins released from 
the anterior pituitary (FSH and LH) were discussed earlier. 


The primary hormone produced by the male testes is testosterone, a steroid 
hormone important in the development of the male reproductive system, the 
maturation of sperm cells, and the development of male secondary sex 
characteristics such as a deepened voice, body hair, and increased muscle 
mass. Interestingly, testosterone is also produced in the female ovaries, but 
at a much reduced level. In addition, the testes produce the peptide hormone 
inhibin, which inhibits the secretion of FSH from the anterior pituitary 
gland. FSH stimulates spermatogenesis. 


The primary hormones produced by the ovaries are estrogens, which 
include estradiol, estriol, and estrone. Estrogens play an important role in a 
larger number of physiological processes, including the development of the 
female reproductive system, regulation of the menstrual cycle, the 
development of female secondary sex characteristics such as increased 
adipose tissue and the development of breast tissue, and the maintenance of 
pregnancy. Another significant ovarian hormone is progesterone, which 
contributes to regulation of the menstrual cycle and is important in 
preparing the body for pregnancy as well as maintaining pregnancy. In 
addition, the granulosa cells of the ovarian follicles produce inhibin, which 
—as in males—inhibits the secretion of FSH.During the initial stages of 
pregnancy, an organ called the placenta develops within the uterus. The 
placenta supplies oxygen and nutrients to the fetus, excretes waste products, 
and produces and secretes estrogens and progesterone. The placenta 
produces human chorionic gonadotropin (hCG) as well. The hCG hormone 
promotes progesterone synthesis and reduces the mother’s immune function 
to protect the fetus from immune rejection. It also secretes human placental 


lactogen (hPL), which plays a role in preparing the breasts for lactation, and 
relaxin, which is thought to help soften and widen the pubic symphysis in 
preparation for childbirth. The hormones controlling reproduction are 


summarized in [link]. 


Reproductive Hormones 


Associated 
Gonad hormones 
Testes Testosterone 
Testes Inhibin 
Estrogens 
Ovaries and 
progesterone 
Human 
Placenta chorionic 
gonadotropin 


Chemical 
class 


Steroid 


Protein 


Steroid 


Protein 


Effect 


Stimulates 
development of male 
secondary sex 
characteristics and 
sperm production 


Inhibits FSH release 
from pituitary 


Stimulate 
development of 
female secondary sex 
characteristics and 
prepare the body for 
childbirth 


Promotes 
progesterone 
synthesis during 
pregnancy and 
inhibits immune 
response against fetus 


Note: 

Everyday Connections 

Anabolic Steroids 

The endocrine system can be exploited for illegal or unethical purposes. A 
prominent example of this is the use of steroid drugs by professional 
athletes. 

Commonly used for performance enhancement, anabolic steroids are 
synthetic versions of the male sex hormone, testosterone. By boosting 
natural levels of this hormone, athletes experience increased muscle mass. 
Synthetic versions of human growth hormone are also used to build muscle 
mass. 

The use of performance-enhancing drugs is banned by all major collegiate 
and professional sports organizations in the United States because they 
impart an unfair advantage to athletes who take them. In addition, the 
drugs can cause significant and dangerous side effects. For example, 
anabolic steroid use can increase cholesterol levels, raise blood pressure, 
and damage the liver. Altered testosterone levels (both too low or too high) 
have been implicated in causing structural damage to the heart, and 
increasing the risk for cardiac arrhythmias, heart attacks, congestive heart 
failure, and sudden death. Paradoxically, steroids can have a feminizing 
effect in males, including shriveled testicles and enlarged breast tissue. In 
females, their use can cause masculinizing effects such as an enlarged 
clitoris and growth of facial hair. In both sexes, their use can promote 
increased aggression (commonly known as “roid-rage”), depression, sleep 
disturbances, severe acne, and infertility. 


Chapter Review 


The male and female reproductive system is regulated by follicle- 
stimulating hormone (FSH) and luteinizing hormone (LH) produced by the 
anterior lobe of the pituitary gland in response to gonadotropin-releasing 
hormone (GnRH) from the hypothalamus. In males, FSH stimulates sperm 
maturation, which is inhibited by the hormone inhibin. The steroid hormone 
testosterone, a type of androgen, is released in response to LH and is 
responsible for the maturation and maintenance of the male reproductive 


system, as well as the development of male secondary sex characteristics. In 
females, FSH promotes egg maturation and LH signals the secretion of the 
female sex hormones, the estrogens and progesterone. Both of these 
hormones are important in the development and maintenance of the female 
reproductive system, as well as maintaining pregnancy. The placenta 
develops during early pregnancy, and secretes several hormones important 
for maintaining the pregnancy. 


Review Questions 


Exercise: 


Problem:The gonads produce what class of hormones? 


a. amine hormones 
b. peptide hormones 
c. steroid hormones 
d. catecholamines 


Solution: 


G 
Exercise: 
Problem: 


The production of FSH by the anterior pituitary is reduced by which 
hormone? 


a. estrogens 

b. progesterone 
c. relaxin 

d. inhibin 


Solution: 


D 
Exercise: 
Problem: 


The function of the placental hormone human placental lactogen (hPL) 
is to 


a. prepare the breasts for lactation 
b. nourish the placenta 

c. regulate the menstrual cycle 

d. all of the above 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem:Compare and contrast the role of estrogens and progesterone. 


Solution: 


Both estrogens and progesterone are steroid hormones produced by the 
ovaries that help regulate the menstrual cycle. Estrogens play an 
important role in the development of the female reproductive tract and 
secondary sex characteristics. They also help maintain pregnancy. 
Progesterone prepares the body for pregnancy and helps maintain 
pregnancy. 


Exercise: 


Problem: 


Describe the role of placental secretion of relaxin in preparation for 
childbirth. 


Solution: 


Relaxin produced by the placenta is thought to soften and widen the 
pubic symphysis. This increases the size of the pelvic outlet, the birth 
canal through which the fetus passes during vaginal childbirth. 


Glossary 


estrogens 
class of predominantly female sex hormones important for the 
development and growth of the female reproductive tract, secondary 
sex characteristics, the female reproductive cycle, and the maintenance 
of pregnancy 


inhibin 
hormone secreted by the male and female gonads that inhibits FSH 
production by the anterior pituitary 


progesterone 
predominantly female sex hormone important in regulating the female 
reproductive cycle and the maintenance of pregnancy 


testosterone 
steroid hormone secreted by the male testes and important in the 
maturation of sperm cells, growth and development of the male 
reproductive system, and the development of male secondary sex 
characteristics 


The Endocrine Pancreas 
By the end of this section, you will be able to: 


e Describe the location and structure of the pancreas, and the 
morphology and function of the pancreatic islets 
¢ Compare and contrast the functions of insulin and glucagon 


The pancreas is a long, slender organ, most of which is located posterior to 
the bottom half of the stomach ([link]). Although it is primarily an exocrine 
gland, secreting a variety of digestive enzymes, the pancreas has an 
endocrine function. Its pancreatic islets—clusters of cells formerly known 
as the islets of Langerhans—secrete the hormones glucagon, insulin, 
somatostatin, and pancreatic polypeptide (PP). 

Pancreas 


Splenic artery 


Pancreatic 
hormones: 
¢ Insulin 
¢ Glucagon 


Spleen 


Pancreatic islets 


Bile duct (from 
gall bladder) 


Common bile duct 


Duodenum of 
small intestine 


Acinar cells 
secrete digestive 
enzymes 


Pancreatic duct 


} ~~ Exocrine acinus 


The pancreatic exocrine function involves the acinar 
cells secreting digestive enzymes that are transported 
into the small intestine by the pancreatic duct. Its 
endocrine function involves the secretion of insulin 
(produced by beta cells) and glucagon (produced by 
alpha cells) within the pancreatic islets. These two 
hormones regulate the rate of glucose metabolism in the 
body. The micrograph reveals pancreatic islets. LM x 
760. (Micrograph provided by the Regents of University 
of Michigan Medical School © 2012) 


Note: 


| 


[=] 


1 


- - 
mss’ OPENStax COLLEGE 


pot 


Data 


View the University of Michigan WebScope to explore the tissue sample in 
greater detail. 


Cells and Secretions of the Pancreatic Islets 


The pancreatic islets each contain four varieties of cells: 


The alpha cell produces the hormone glucagon and makes up 
approximately 20 percent of each islet. Glucagon plays an important 
role in blood glucose regulation; low blood glucose levels stimulate its 
release. 

The beta cell produces the hormone insulin and makes up 
approximately 75 percent of each islet. Elevated blood glucose levels 
stimulate the release of insulin. 

The delta cell accounts for four percent of the islet cells and secretes 
the peptide hormone somatostatin. Recall that somatostatin is also 
released by the hypothalamus (as GHIH), and the stomach and 
intestines also secrete it. An inhibiting hormone, pancreatic 
somatostatin inhibits the release of both glucagon and insulin. 

The PP cell accounts for about one percent of islet cells and secretes 
the pancreatic polypeptide hormone. It is thought to play a role in 
appetite, as well as in the regulation of pancreatic exocrine and 
endocrine secretions. Pancreatic polypeptide released following a meal 


may reduce further food consumption; however, it is also released in 
response to fasting. 


Regulation of Blood Glucose Levels by Insulin and Glucagon 


Glucose is required for cellular respiration and is the preferred fuel for all 
body cells. The body derives glucose from the breakdown of the 
carbohydrate-containing foods and drinks we consume. Glucose not 
immediately taken up by cells for fuel can be stored by the liver and 
muscles as glycogen, or converted to triglycerides and stored in the adipose 
tissue. Hormones regulate both the storage and the utilization of glucose as 
required. Receptors located in the pancreas sense blood glucose levels, and 
subsequently the pancreatic cells secrete glucagon or insulin to maintain 
normal levels. 


Glucagon 


Receptors in the pancreas can sense the decline in blood glucose levels, 
such as during periods of fasting or during prolonged labor or exercise 
({link]). In response, the alpha cells of the pancreas secrete the hormone 
glucagon, which has several effects: 


e It stimulates the liver to convert its stores of glycogen back into 
glucose. This response is known as glycogenolysis. The glucose is 
then released into the circulation for use by body cells. 

e It stimulates the liver to take up amino acids from the blood and 
convert them into glucose. This response is known as gluconeogenesis. 

e It stimulates lipolysis, the breakdown of stored triglycerides into free 
fatty acids and glycerol. Some of the free glycerol released into the 
bloodstream travels to the liver, which converts it into glucose. This is 
also a form of gluconeogenesis. 


Taken together, these actions increase blood glucose levels. The activity of 
glucagon is regulated through a negative feedback mechanism; rising blood 
glucose levels inhibit further glucagon production and secretion. 
Homeostatic Regulation of Blood Glucose Levels 


Insulin release: Insulin effects: 


¢ Beta cells of pancreas 
release insulin * Triggers body cells to take up 
glucose from the blood and 
utilize it in cellular respiration 


Splenic artery 


¢ Inhibits glycogenolysis 
— glucose is removed from the 
blood and stored as glycogen 
in the liver 


¢ Inhibits gluconeogenesis Rough ER 
- amino acids and free glycerol 
are NOT converted to 


glucose in the ER 
Smooth ER 


Blood glucose 
concentration 
decreases 


Hyperglycemia 

(elevated blood glucose) 

Hypoglycemia 

(low blood glucose) yh 


START: Homeostasis 


(70-110 mg/dL) ee 


Blood glucose 


concentration 
increases 


Glucagon effects: 


Inhibits body cells from taking 
up glucose from the blood and 
Glucagon release: utilizing it in cellular respiration 
¢ Alpha cells of pancreas 
| | 
nee ee Splenic artery * Stimulates glycogenolysis 
— glycogen in the liver is 
broken down into glucose 
and released into the blood 


¢ Stimulates gluconeogenesis 
— amino acids and free glycerol Rough ER 
are converted to glucose in 


the ER and released into the 
blood 
Smooth ER 


Blood glucose concentration is tightly maintained between 70 
mg/dL and 110 mg/dL. If blood glucose concentration rises 


above this range, insulin is released, which stimulates body 
cells to remove glucose from the blood. If blood glucose 
concentration drops below this range, glucagon is released, 
which stimulates body cells to release glucose into the blood. 


Insulin 


The primary function of insulin is to facilitate the uptake of glucose into 
body cells. Red blood cells, as well as cells of the brain, liver, kidneys, and 
the lining of the small intestine, do not have insulin receptors on their cell 
membranes and do not require insulin for glucose uptake. Although all 
other body cells do require insulin if they are to take glucose from the 
bloodstream, skeletal muscle cells and adipose cells are the primary targets 
of insulin. 


The presence of food in the intestine triggers the release of gastrointestinal 
tract hormones such as glucose-dependent insulinotropic peptide 
(previously known as gastric inhibitory peptide). This is in turn the initial 
trigger for insulin production and secretion by the beta cells of the pancreas. 
Once nutrient absorption occurs, the resulting surge in blood glucose levels 
further stimulates insulin secretion. 


Precisely how insulin facilitates glucose uptake is not entirely clear. 
However, insulin appears to activate a tyrosine kinase receptor, triggering 
the phosphorylation of many substrates within the cell. These multiple 
biochemical reactions converge to support the movement of intracellular 
vesicles containing facilitative glucose transporters to the cell membrane. In 
the absence of insulin, these transport proteins are normally recycled slowly 
between the cell membrane and cell interior. Insulin triggers the rapid 
movement of a pool of glucose transporter vesicles to the cell membrane, 
where they fuse and expose the glucose transporters to the extracellular 
fluid. The transporters then move glucose by facilitated diffusion into the 
cell interior. 


Visit this link to view an animation describing the location and function of 
the pancreas. What goes wrong in the function of insulin in type 2 
diabetes? 


Insulin also reduces blood glucose levels by stimulating glycolysis, the 
metabolism of glucose for generation of ATP. Moreover, it stimulates the 
liver to convert excess glucose into glycogen for storage, and it inhibits 
enzymes involved in glycogenolysis and gluconeogenesis. Finally, insulin 
promotes triglyceride and protein synthesis. The secretion of insulin is 
regulated through a negative feedback mechanism. As blood glucose levels 
decrease, further insulin release is inhibited. The pancreatic hormones are 
summarized in [link]. 


Hormones of the Pancreas 


Chemical 
Associated hormones class Effect 
Insulin (beta cells) Protein Reduces blood glucose 


levels 


Hormones of the Pancreas 


Chemical 
Associated hormones class Effect 
Glucagon (alpha cells) Protein Tacreases DiOOG eileos: 
levels 
Somatostatin (delta Peavieia Inhibits insulin and 
cells) glucagon release 
Palicrealle/poly pepude Protein Role in appetite 
(PP cells) 
Note: 


Disorders of the... 

Endocrine System: Diabetes Mellitus 

Dysfunction of insulin production and secretion, as well as the target cells’ 
responsiveness to insulin, can lead to a condition called diabetes mellitus. 
An increasingly common disease, diabetes mellitus has been diagnosed in 
more than 18 million adults in the United States, and more than 200,000 
children. It is estimated that up to 7 million more adults have the condition 
but have not been diagnosed. In addition, approximately 79 million people 
in the US are estimated to have pre-diabetes, a condition in which blood 
glucose levels are abnormally high, but not yet high enough to be classified 
as diabetes. 

There are two main forms of diabetes mellitus. Type 1 diabetes is an 
autoimmune disease affecting the beta cells of the pancreas. Certain genes 
are recognized to increase susceptibility. The beta cells of people with type 
1 diabetes do not produce insulin; thus, synthetic insulin must be 
administered by injection or infusion. This form of diabetes accounts for 
less than five percent of all diabetes cases. 

Type 2 diabetes accounts for approximately 95 percent of all cases. It is 
acquired, and lifestyle factors such as poor diet, inactivity, and the presence 


of pre-diabetes greatly increase a person’s risk. About 80 to 90 percent of 
people with type 2 diabetes are overweight or obese. In type 2 diabetes, 
cells become resistant to the effects of insulin. In response, the pancreas 
increases its insulin secretion, but over time, the beta cells become 
exhausted. In many cases, type 2 diabetes can be reversed by moderate 
weight loss, regular physical activity, and consumption of a healthy diet; 
however, if blood glucose levels cannot be controlled, the diabetic will 
eventually require insulin. 

Two of the early manifestations of diabetes are excessive urination and 
excessive thirst. They demonstrate how the out-of-control levels of glucose 
in the blood affect kidney function. The kidneys are responsible for 
filtering glucose from the blood. Excessive blood glucose draws water into 
the urine, and as a result the person eliminates an abnormally large quantity 
of sweet urine. The use of body water to dilute the urine leaves the body 
dehydrated, and so the person is unusually and continually thirsty. The 
person may also experience persistent hunger because the body cells are 
unable to access the glucose in the bloodstream. 

Over time, persistently high levels of glucose in the blood injure tissues 
throughout the body, especially those of the blood vessels and nerves. 
Inflammation and injury of the lining of arteries lead to atherosclerosis and 
an increased risk of heart attack and stroke. Damage to the microscopic 
blood vessels of the kidney impairs kidney function and can lead to kidney 
failure. Damage to blood vessels that serve the eyes can lead to blindness. 
Blood vessel damage also reduces circulation to the limbs, whereas nerve 
damage leads to a loss of sensation, called neuropathy, particularly in the 
hands and feet. Together, these changes increase the risk of injury, 
infection, and tissue death (necrosis), contributing to a high rate of toe, 
foot, and lower leg amputations in people with diabetes. Uncontrolled 
diabetes can also lead to a dangerous form of metabolic acidosis called 
ketoacidosis. Deprived of glucose, cells increasingly rely on fat stores for 
fuel. However, in a glucose-deficient state, the liver is forced to use an 
alternative lipid metabolism pathway that results in the increased 
production of ketone bodies (or ketones), which are acidic. The build-up of 
ketones in the blood causes ketoacidosis, which—if left untreated—may 
lead to a life-threatening “diabetic coma.” Together, these complications 
make diabetes the seventh leading cause of death in the United States. 


Diabetes is diagnosed when lab tests reveal that blood glucose levels are 
higher than normal, a condition called hyperglycemia. The treatment of 
diabetes depends on the type, the severity of the condition, and the ability 
of the patient to make lifestyle changes. As noted earlier, moderate weight 
loss, regular physical activity, and consumption of a healthful diet can 
reduce blood glucose levels. Some patients with type 2 diabetes may be 
unable to control their disease with these lifestyle changes, and will require 
medication. Historically, the first-line treatment of type 2 diabetes was 
insulin. Research advances have resulted in alternative options, including 
medications that enhance pancreatic function. 


Note: 


[=] [= 


— 
meee, OPENStAX COLLEGE 
Aor 


= 


Visit this link to view an animation describing the role of insulin and the 
pancreas in diabetes. 


Chapter Review 


The pancreas has both exocrine and endocrine functions. The pancreatic 
islet cell types include alpha cells, which produce glucagon; beta cells, 
which produce insulin; delta cells, which produce somatostatin; and PP 
cells, which produce pancreatic polypeptide. Insulin and glucagon are 
involved in the regulation of glucose metabolism. Insulin is produced by the 
beta cells in response to high blood glucose levels. It enhances glucose 
uptake and utilization by target cells, as well as the storage of excess 
glucose for later use. Dysfunction of the production of insulin or target cell 
resistance to the effects of insulin causes diabetes mellitus, a disorder 


characterized by high blood glucose levels. The hormone glucagon is 
produced and secreted by the alpha cells of the pancreas in response to low 
blood glucose levels. Glucagon stimulates mechanisms that increase blood 
glucose levels, such as the catabolism of glycogen into glucose. 


Interactive Link Questions 


Exercise: 


Problem: 


Visit this link to view an animation describing the location and 
function of the pancreas. What goes wrong in the function of insulin in 
type 2 diabetes? 


Solution: 


Insulin is overproduced. 


Review Questions 


Exercise: 


Problem: 


If an autoimmune disorder targets the alpha cells, production of which 
hormone would be directly affected? 


a. somatostatin 

b. pancreatic polypeptide 
c. insulin 

d. glucagon 


Solution: 


D 


Exercise: 


Problem: Which of the following statements about insulin is true? 


a. Insulin acts as a transport protein, carrying glucose across the cell 
membrane. 

b. Insulin facilitates the movement of intracellular glucose 
transporters to the cell membrane. 

c. Insulin stimulates the breakdown of stored glycogen into glucose. 

d. Insulin stimulates the kidneys to reabsorb glucose into the 
bloodstream. 


Solution: 


B 


Critical Thinking Questions 


Exercise: 


Problem: 


What would be the physiological consequence of a disease that 
destroyed the beta cells of the pancreas? 


Solution: 


The beta cells produce the hormone insulin, which is important in the 
regulation of blood glucose levels. All insulin-dependent cells of the 
body require insulin in order to take up glucose from the bloodstream. 
Destruction of the beta cells would result in an inability to produce and 
secrete insulin, leading to abnormally high blood glucose levels and 
the disease called type 1 diabetes mellitus. 


Exercise: 


Problem: 


Why is foot care extremely important for people with diabetes 
mellitus? 


Solution: 


Excessive blood glucose levels damage the blood vessels and nerves of 
the body’s extremities, increasing the risk for injury, infection, and 
tissue death. Loss of sensation to the feet means that a diabetic patient 
will not be able to feel foot trauma, such as from ill-fitting shoes. Even 
minor injuries commonly lead to infection, which , can progress to 
tissue death without proper care, requiring amputation. 


Glossary 


alpha cell 
pancreatic islet cell type that produces the hormone glucagon 


beta cell 
pancreatic islet cell type that produces the hormone insulin 


delta cell 
minor cell type in the pancreas that secretes the hormone somatostatin 


diabetes mellitus 
condition caused by destruction or dysfunction of the beta cells of the 
pancreas or cellular resistance to insulin that results in abnormally high 
blood glucose levels 


glucagon 
pancreatic hormone that stimulates the catabolism of glycogen to 
glucose, thereby increasing blood glucose levels 


hyperglycemia 
abnormally high blood glucose levels 


insulin 
pancreatic hormone that enhances the cellular uptake and utilization of 
glucose, thereby decreasing blood glucose levels 


pancreas 
organ with both exocrine and endocrine functions located posterior to 
the stomach that is important for digestion and the regulation of blood 
glucose 


pancreatic islets 
specialized clusters of pancreatic cells that have endocrine functions; 
also called islets of Langerhans 


PP cell 
minor cell type in the pancreas that secretes the hormone pancreatic 
polypeptide 


Organs with Secondary Endocrine Functions 
By the end of this section, you will be able to: 


e Identify the organs with a secondary endocrine function, the hormone 
they produce, and its effects 


In your study of anatomy and physiology, you have already encountered a 
few of the many organs of the body that have secondary endocrine 
functions. Here, you will learn about the hormone-producing activities of 
the heart, gastrointestinal tract, kidneys, skeleton, adipose tissue, skin, and 
thymus. 


Heart 


When the body experiences an increase in blood volume or pressure, the 
cells of the heart’s atrial wall stretch. In response, specialized cells in the 
wall of the atria produce and secrete the peptide hormone atrial natriuretic 
peptide (ANP). ANP signals the kidneys to reduce sodium reabsorption, 
thereby decreasing the amount of water reabsorbed from the urine filtrate 
and reducing blood volume. Other actions of ANP include the inhibition of 
renin secretion and the initiation of the renin-angiotensin-aldosterone 
system (RAAS) and vasodilation. Therefore, ANP aids in decreasing blood 
pressure, blood volume, and blood sodium levels. 


Gastrointestinal Tract 


The endocrine cells of the GI tract are located in the mucosa of the stomach 
and small intestine. Some of these hormones are secreted in response to 
eating a meal and aid in digestion. An example of a hormone secreted by 
the stomach cells is gastrin, a peptide hormone secreted in response to 
stomach distention that stimulates the release of hydrochloric acid. Secretin 
is a peptide hormone secreted by the small intestine as acidic chyme 
(partially digested food and fluid) moves from the stomach. It stimulates the 
release of bicarbonate from the pancreas, which buffers the acidic chyme, 
and inhibits the further secretion of hydrochloric acid by the stomach. 
Cholecystokinin (CCK) is another peptide hormone released from the small 
intestine. It promotes the secretion of pancreatic enzymes and the release of 


bile from the gallbladder, both of which facilitate digestion. Other 
hormones produced by the intestinal cells aid in glucose metabolism, such 
as by stimulating the pancreatic beta cells to secrete insulin, reducing 
glucagon secretion from the alpha cells, or enhancing cellular sensitivity to 
insulin. 


Kidneys 


The kidneys participate in several complex endocrine pathways and 
produce certain hormones. A decline in blood flow to the kidneys 
stimulates them to release the enzyme renin, triggering the renin- 
angiotensin-aldosterone (RAAS) system, and stimulating the reabsorption 
of sodium and water. The reabsorption increases blood flow and blood 
pressure. The kidneys also play a role in regulating blood calcium levels 
through the production of calcitriol from vitamin D3, which is released in 
response to the secretion of parathyroid hormone (PTH). In addition, the 
kidneys produce the hormone erythropoietin (EPO) in response to low 
oxygen levels. EPO stimulates the production of red blood cells 
(erythrocytes) in the bone marrow, thereby increasing oxygen delivery to 
tissues. You may have heard of EPO as a performance-enhancing drug (in a 
synthetic form). 


Skeleton 


Although bone has long been recognized as a target for hormones, only 
recently have researchers recognized that the skeleton itself produces at 
least two hormones. Fibroblast growth factor 23 (FGF23) is produced by 
bone cells in response to increased blood levels of vitamin D3 or phosphate. 
It triggers the kidneys to inhibit the formation of calcitriol from vitamin D3 
and to increase phosphorus excretion. Osteocalcin, produced by osteoblasts, 
stimulates the pancreatic beta cells to increase insulin production. It also 
acts on peripheral tissues to increase their sensitivity to insulin and their 
utilization of glucose. 


Adipose Tissue 


Adipose tissue produces and secretes several hormones involved in lipid 
metabolism and storage. One important example is leptin, a protein 
manufactured by adipose cells that circulates in amounts directly 
proportional to levels of body fat. Leptin is released in response to food 
consumption and acts by binding to brain neurons involved in energy intake 
and expenditure. Binding of leptin produces a feeling of satiety after a meal, 
thereby reducing appetite. It also appears that the binding of leptin to brain 
receptors triggers the sympathetic nervous system to regulate bone 
metabolism, increasing deposition of cortical bone. Adiponectin—another 
hormone synthesized by adipose cells—appears to reduce cellular insulin 
resistance and to protect blood vessels from inflammation and 
atherosclerosis. Its levels are lower in people who are obese, and rise 
following weight loss. 


Skin 


The skin functions as an endocrine organ in the production of the inactive 
form of vitamin D3, cholecalciferol. When cholesterol present in the 
epidermis is exposed to ultraviolet radiation, it is converted to 
cholecalciferol, which then enters the blood. In the liver, cholecalciferol is 
converted to an intermediate that travels to the kidneys and is further 
converted to calcitriol, the active form of vitamin D3. Vitamin D is 
important in a variety of physiological processes, including intestinal 
calcium absorption and immune system function. In some studies, low 
levels of vitamin D have been associated with increased risks of cancer, 
severe asthma, and multiple sclerosis. Vitamin D deficiency in children 
causes rickets, and in adults, osteomalacia—both of which are characterized 
by bone deterioration. 


Thymus 


The thymus is an organ of the immune system that is larger and more 
active during infancy and early childhood, and begins to atrophy as we age. 
Its endocrine function is the production of a group of hormones called 
thymosins that contribute to the development and differentiation of T 
lymphocytes, which are immune cells. Although the role of thymosins is 


not yet well understood, it is clear that they contribute to the immune 
response. Thymosins have been found in tissues other than the thymus and 
have a wide variety of functions, so the thymosins cannot be strictly 
categorized as thymic hormones. 


Liver 


The liver is responsible for secreting at least four important hormones or 
hormone precursors: insulin-like growth factor (somatomedin), 
angiotensinogen, thrombopoetin, and hepcidin. Insulin-like growth factor-1 
is the immediate stimulus for growth in the body, especially of the bones. 
Angiotensinogen is the precursor to angiotensin, mentioned earlier, which 
increases blood pressure. Thrombopoetin stimulates the production of the 
blood’s platelets. Hepcidins block the release of iron from cells in the body, 
helping to regulate iron homeostasis in our body fluids. The major 
hormones of these other organs are summarized in [Link]. 


Organs with Secondary Endocrine Functions and Their Major 
Hormones 


Organ Major hormones Effects 
Reduces blood 
Heart Atrial natriuretic volume, blood 
peptide (ANP) pressure, and Na* 
concentration 
Gastrointestinal Gastrin, secretin, and a digestion oad 
ae and buffering of 
tract cholecystokinin 


stomach acids 


Organs with Secondary Endocrine Functions and Their Major 


Hormones 


Organ 


Gastrointestinal 
tract 


Kidneys 


Kidneys 


Kidneys 


Skeleton 


Skeleton 


Adipose tissue 


Adipose tissue 


Major hormones 


Glucose-dependent 
insulinotropic peptide 
(GIP) and glucagon- 
like peptide 1 (GLP- 
1) 


Renin 


Calcitriol 


Erythropoietin 


FGF23 


Osteocalcin 


Leptin 


Adiponectin 


Effects 


Stimulate beta cells 
of the pancreas to 
release insulin 


Stimulates release of 
aldosterone 


Aids in the 
absorption of Ca** 


Triggers the 
formation of red 
blood cells in the 
bone marrow 


Inhibits production of 
calcitriol and 
increases phosphate 
excretion 


Increases insulin 
production 


Promotes satiety 
signals in the brain 


Reduces insulin 
resistance 


Organs with Secondary Endocrine Functions and Their Major 


Hormones 


Organ 


Skin 


Thymus (and 
other organs) 


Liver 
Liver 


Liver 


Liver 


Chapter Review 


Major hormones 


Cholecalciferol 


Thymosins 


Insulin-like growth 
factor-1 


Angiotensinogen 


Thrombopoetin 


Hepcidin 


Effects 


Modified to form 
vitamin D 


Among other things, 
aids in the 
development of T 
lymphocytes of the 
immune system 


Stimulates bodily 
growth 


Raises blood pressure 


Causes increase in 
platelets 


Blocks release of iron 
into body fluids 


Some organs have a secondary endocrine function. For example, the walls 
of the atria of the heart produce the hormone atrial natriuretic peptide 
(ANP), the gastrointestinal tract produces the hormones gastrin, secretin, 
and cholecystokinin, which aid in digestion, and the kidneys produce 
erythropoietin (EPO), which stimulates the formation of red blood cells. 
Even bone, adipose tissue, and the skin have secondary endocrine functions. 


Review Questions 


Exercise: 


Problem:The walls of the atria produce which hormone? 


a. cholecystokinin 

b. atrial natriuretic peptide 
c. renin 

d. calcitriol 


Solution: 


B 


Exercise: 


Problem:The end result of the RAAS is to 


a. reduce blood volume 
b. increase blood glucose 
c. reduce blood pressure 
d. increase blood pressure 


Solution: 


D 


Exercise: 


Problem: Athletes may take synthetic EPO to boost their 


a. blood calcium levels 

b. secretion of growth hormone 
c. blood oxygen levels 

d. muscle mass 


Solution: 


C 
Exercise: 


Problem: 
Hormones produced by the thymus play a role in the 


a. development of T cells 

b. preparation of the body for childbirth 

c. regulation of appetite 

d. release of hydrochloric acid in the stomach 


Solution: 


A 


Critical Thinking Questions 


Exercise: 


Problem:Summarize the role of GI tract hormones following a meal. 


Solution: 


The presence of food in the GI tract stimulates the release of hormones 
that aid in digestion. For example, gastrin is secreted in response to 
stomach distention and causes the release of hydrochloric acid in the 
stomach. Secretin is secreted when acidic chyme enters the small 
intestine, and stimulates the release of pancreatic bicarbonate. In the 
presence of fat and protein in the duodenum, CCK stimulates the 
release of pancreatic digestive enzymes and bile from the gallbladder. 
Other GI tract hormones aid in glucose metabolism and other 
functions. 


Exercise: 


Problem: 


Compare and contrast the thymus gland in infancy and adulthood. 


Solution: 


The thymus gland is important for the development and maturation of 
T cells. During infancy and early childhood, the thymus gland is large 
and very active, as the immune system is still developing. During 
adulthood, the thymus gland atrophies because the immune system is 
already developed. 


Glossary 


atrial natriuretic peptide (ANP) 
peptide hormone produced by the walls of the atria in response to high 
blood pressure, blood volume, or blood sodium that reduces the 
reabsorption of sodium and water in the kidneys and promotes 
vasodilation 


erythropoietin (EPO) 
protein hormone secreted in response to low oxygen levels that 
triggers the bone marrow to produce red blood cells 


leptin 
protein hormone secreted by adipose tissues in response to food 
consumption that promotes satiety 


thymosins 
hormones produced and secreted by the thymus that play an important 
role in the development and differentiation of T cells 


thymus 
organ that is involved in the development and maturation of T-cells 
and is particularly active during infancy and childhood 


Male Anatomy 
By the end of this section, you will be able to: 


e Describe the structure and function of the organs of the male 
reproductive system 

¢ Describe the structure and function of the sperm cell 

e Explain the events during spermatogenesis that produce haploid sperm 
from diploid cells 

e Identify the importance of testosterone in male reproductive function 


Unique for its role in human reproduction, a gamete is a specialized sex 
cell carrying 23 chromosomes—one half the number in body cells. At 
fertilization, the chromosomes in one male gamete, called a sperm (or 
spermatozoon), combine with the chromosomes in one female gamete, 
called an oocyte. The function of the male reproductive system ((link]) is to 
produce sperm and transfer them to the female reproductive tract. The 
paired testes are a crucial component in this process, as they produce both 
sperm and androgens, the hormones that support male reproductive 
physiology. In humans, the most important male androgen is testosterone. 
Several accessory organs and ducts aid the process of sperm maturation and 
transport the sperm and other seminal components to the penis, which 
delivers sperm to the female reproductive tract. In this section, we examine 
each of these different structures, and discuss the process of sperm 
production and transport. 

Male Reproductive System 


(a) Uncircumcised penis (b) Circumcised penis 


Penis 


Prepuce 
(foreskin) 


(c) Male Reproductive System: lateral view 


i 


Ductus (vas) deferens 
Suspensory ligament of penis 


Ampulla of ductus deferens 


a : aml | a) | Seminal vesicle 
Pubic symphysis ME; if 
Ejaculatory duct 
Prostatic urethra 
Deep muscles of perineum 
Bulbourethral gland 
ie Muscles of perineum 
Corpus cavernosum —— } surrounding anus 
= Membranous urethra 
Spongy urethra 
Testis 
Epididymis 
Corpus spongiosum Scrotum 


External urethral opening 


The structures of the male reproductive system include the 
testes, the epididymides, the penis, and the ducts and glands 
that produce and carry semen. Sperm exit the scrotum through 
the ductus deferens, which is bundled in the spermatic cord. 
The seminal vesicles and prostate gland add fluids to the sperm 
to create semen. 


Scrotum 


The testes are located in a skin-covered, highly pigmented, muscular sack 
called the scrotum that extends from the body behind the penis (see [Link]). 
This location is important in sperm production, which occurs within the 
testes, and proceeds more efficiently when the testes are kept 2 to 4°C 
below core body temperature. 


The dartos muscle makes up the subcutaneous muscle layer of the scrotum 
({link]). It continues internally to make up the scrotal septum, a wall that 
divides the scrotum into two compartments, each housing one testis. 
Descending from the internal oblique muscle of the abdominal wall are the 
two cremaster muscles, which cover each testis like a muscular net. By 
contracting simultaneously, the dartos and cremaster muscles can elevate 
the testes in cold weather (or water), moving the testes closer to the body 
and decreasing the surface area of the scrotum to retain heat. Alternatively, 
as the environmental temperature increases, the scrotum relaxes, moving 
the testes farther from the body core and increasing scrotal surface area, 
which promotes heat loss. Externally, the scrotum has a raised medial 
thickening on the surface called the raphae. 
The Scrotum and Testes 

External view of scrotum Muscle layer Deep tissues 


Ductus Spermatic 
deferens cord 


Testicular 
artery 


Autonomic 
nerve 


Lymphatic 
vessel 


|> Testis 


Y 
Cremaster 
muscles muscles 


This anterior view shows the structures of the scrotum and 
testes. 


Testes 


The testes (singular = testis) are the male gonads—that is, the male 
reproductive organs. They produce both sperm and androgens, such as 
testosterone, and are active throughout the reproductive lifespan of the 
male. 


Paired ovals, the testes are each approximately 4 to 5 cm in length and are 
housed within the scrotum (see [link]). They are surrounded by two distinct 
layers of protective connective tissue ({link]). The outer tunica vaginalis is a 
serous membrane that has both a parietal and a thin visceral layer. Beneath 
the tunica vaginalis is the tunica albuginea, a tough, white, dense 
connective tissue layer covering the testis itself. Not only does the tunica 
albuginea cover the outside of the testis, it also invaginates to form septa 
that divide the testis into 300 to 400 structures called lobules. Within the 
lobules, sperm develop in structures called seminiferous tubules. During the 
seventh month of the developmental period of a male fetus, each testis 
moves through the abdominal musculature to descend into the scrotal 
cavity. This is called the “descent of the testis.” Cryptorchidism is the 
clinical term used when one or both of the testes fail to descend into the 
scrotum prior to birth. 

Anatomy of the Testis 


} Into inguinal canal 


Spermatic cord __a 
Cremaster 
muscle 
Efferent Tunica vaginalis 
ductule 
Body of Head of» 
epididymis / epididymis 
Ductus | Seminiferous 
deferens tubule lobules 
' nC = Septa (tunica 
Rete w4 FP BOD oars albuginea) 
testis ——s * “4 
Tunica albuginea 
Straight 
tubule 


Tail of epididymis 


This sagittal view shows the seminiferous tubules, 
the site of sperm production. Formed sperm are 
transferred to the epididymis, where they mature. 
They leave the epididymis during an ejaculation 
via the ductus deferens. 


The tightly coiled seminiferous tubules form the bulk of each testis. They 
are composed of developing sperm cells surrounding a lumen, the hollow 
center of the tubule, where formed sperm are released into the duct system 
of the testis. Specifically, from the lumens of the seminiferous tubules, 
sperm move into the straight tubules (or tubuli recti), and from there into a 
fine meshwork of tubules called the rete testes. Sperm leave the rete testes, 
and the testis itself, through the 15 to 20 efferent ductules that cross the 
tunica albuginea. 


Inside the seminiferous tubules are six different cell types. These include 
supporting cells called sustentacular cells, as well as five types of 
developing sperm cells called germ cells. Germ cell development 
progresses from the basement membrane—at the perimeter of the tubule— 
toward the lumen. Let’s look more closely at these cell types. 


Sertoli Cells 


Surrounding all stages of the developing sperm cells are elongate, 
branching Sertoli cells. Sertoli cells are a type of supporting cell called a 
sustentacular cell, or sustentocyte, that are typically found in epithelial 
tissue. Sertoli cells secrete signaling molecules that promote sperm 
production and can control whether germ cells live or die. They extend 
physically around the germ cells from the peripheral basement membrane 
of the seminiferous tubules to the lumen. Tight junctions between these 
sustentacular cells create the blood—testis barrier, which keeps bloodborne 
substances from reaching the germ cells and, at the same time, keeps 
surface antigens on developing germ cells from escaping into the 
bloodstream and prompting an autoimmune response. 


Germ Cells 


The least mature cells, the spermatogonia (singular = spermatogonium), 
line the basement membrane inside the tubule. Spermatogonia are the stem 
cells of the testis, which means that they are still able to differentiate into a 
variety of different cell types throughout adulthood. Spermatogonia divide 
to produce primary and secondary spermatocytes, then spermatids, which 
finally produce formed sperm. The process that begins with spermatogonia 
and concludes with the production of sperm is called spermatogenesis. 


Spermatogenesis 


As just noted, spermatogenesis occurs in the seminiferous tubules that form 
the bulk of each testis (see [link]). The process begins at puberty, after 
which time sperm are produced constantly throughout a man’s life. One 
production cycle, from spermatogonia through formed sperm, takes 
approximately 64 days. A new cycle starts approximately every 16 days, 
although this timing is not synchronous across the seminiferous tubules. 
Sperm counts—the total number of sperm a man produces—slowly decline 
after age 35, and some studies suggest that smoking can lower sperm counts 
irrespective of age. 


The process of spermatogenesis begins with mitosis of the diploid 
spermatogonia ([link]). Because these cells are diploid (2n), they each have 
a complete copy of the father’s genetic material, or 46 chromosomes. 
However, mature gametes are haploid (1n), containing 23 chromosomes— 
meaning that daughter cells of spermatogonia must undergo a second 
cellular division through the process of meiosis. 

Spermatogenesis 


(a) Spermatogenesis 


foe 


'2n ) | 2n ) Primary spermatocyte 


BU interstitial 


=— Lymphatic 
capillary 


\)S Sertoli 
}| (sustentacular) 
cell 


a Early 
G spermatids 


be 


S ig <z 5. ss a ; 
Ge Ga Ga Primary WA Arteriole 
in in n) (1) spermatid i ie /e 


ry 


6 re 
| Spermiogenesis 7 ' 
(interstitial) cells 
Spermatozoa — peritubular 
(sperm) capillary 


(a) Mitosis of a spermatogonial stem cell involves a single cell 
division that results in two identical, diploid daughter cells 


(spermatogonia to primary spermatocyte). Meiosis has two 
rounds of cell division: primary spermatocyte to secondary 
spermatocyte, and then secondary spermatocyte to spermatid. 
This produces four haploid daughter cells (spermatids). (b) In 
this electron micrograph of a cross-section of a seminiferous 
tubule from a rat, the lumen is the light-shaded area in the 
center of the image. The location of the primary spermatocytes 
is near the basement membrane, and the early spermatids are 
approaching the lumen (tissue source: rat). EM x 900. 
(Micrograph provided by the Regents of University of 
Michigan Medical School © 2012) 


Two identical diploid cells result from spermatogonia mitosis. One of these 
cells remains a spermatogonium, and the other becomes a primary 
spermatocyte, the next stage in the process of spermatogenesis. As in 
mitosis, DNA is replicated in a primary spermatocyte, before it undergoes a 
cell division called meiosis I. During meiosis I each of the 23 pairs of 
chromosomes separates. This results in two cells, called secondary 
spermatocytes, each with only half the number of chromosomes. Now a 
second round of cell division (meiosis IT) occurs in both of the secondary 
spermatocytes. During meiosis II each of the 23 replicated chromosomes 
divides, similar to what happens during mitosis. Thus, meiosis results in 
separating the chromosome pairs. This second meiotic division results in a 
total of four cells with only half of the number of chromosomes. Each of 
these new cells is a spermatid. Although haploid, early spermatids look 
very similar to cells in the earlier stages of spermatogenesis, with a round 
shape, central nucleus, and large amount of cytoplasm. A process called 
spermiogenesis transforms these early spermatids, reducing the cytoplasm, 
and beginning the formation of the parts of a true sperm. The fifth stage of 
germ cell formation—spermatozoa, or formed sperm—is the end result of 
this process, which occurs in the portion of the tubule nearest the lumen. 
Eventually, the sperm are released into the lumen and are moved along a 
series of ducts in the testis toward a structure called the epididymis for the 
next step of sperm maturation. 


Structure of Formed Sperm 


Sperm are smaller than most cells in the body; in fact, the volume of a 
sperm cell is 85,000 times less than that of the female gamete. 
Approximately 100 to 300 million sperm are produced each day, whereas 
women typically ovulate only one oocyte per month. As is true for most 
cells in the body, the structure of sperm cells speaks to their function. 
Sperm have a distinctive head, mid-piece, and tail region ([link]). The head 
of the sperm contains the extremely compact haploid nucleus with very 
little cytoplasm. These qualities contribute to the overall small size of the 
sperm (the head is only 5 pm long). A structure called the acrosome covers 
most of the head of the sperm cell as a “cap” that is filled with lysosomal 
enzymes important for preparing sperm to participate in fertilization. 
Tightly packed mitochondria fill the mid-piece of the sperm. ATP produced 
by these mitochondria will power the flagellum, which extends from the 
neck and the mid-piece through the tail of the sperm, enabling it to move 
the entire sperm cell. The central strand of the flagellum, the axial filament, 
is formed from one centriole inside the maturing sperm cell during the final 
stages of spermatogenesis. 

Structure of Sperm 


Acrosome Axial filament 
Plasma membrane 


Nucleus Mitochondria 
Centriole 


Flagellum 


Head Mid-piece Tail End 


Sperm cells are divided into a head, containing DNA; a mid- 
piece, containing mitochondria; and a tail, providing motility. 
The acrosome is oval and somewhat flattened. 


Sperm Transport 


To fertilize an egg, sperm must be moved from the seminiferous tubules in 
the testes, through the epididymis, and—later during ejaculation—along the 
length of the penis and out into the female reproductive tract. 


Role of the Epididymis 


From the lumen of the seminiferous tubules, the immotile sperm are 
surrounded by testicular fluid and moved to the epididymis (plural = 
epididymides), a coiled tube attached to the testis where newly formed 
sperm continue to mature (see [link]). Though the epididymis does not take 
up much room in its tightly coiled state, it would be approximately 6 m (20 
feet) long if straightened. It takes an average of 12 days for sperm to move 
through the coils of the epididymis, with the shortest recorded transit time 
in humans being one day. Sperm enter the head of the epididymis and are 
moved along predominantly by the contraction of smooth muscles lining 
the epididymal tubes. As they are moved along the length of the 
epididymis, the sperm further mature and acquire the ability to move under 
their own power. Once inside the female reproductive tract, they will use 
this ability to move independently toward the unfertilized egg. The more 
mature sperm are then stored in the tail of the epididymis (the final section) 
until ejaculation occurs. 


Duct System 


During ejaculation, sperm exit the tail of the epididymis and are pushed by 
smooth muscle contraction to the ductus deferens (also called the vas 
deferens). The ductus deferens is a thick, muscular tube that is bundled 
together inside the scrotum with connective tissue, blood vessels, and 
nerves into a structure called the spermatic cord (see [link] and [link]). 
Because the ductus deferens is physically accessible within the scrotum, 
surgical sterilization to interrupt sperm delivery can be performed by 
cutting and sealing a small section of the ductus (vas) deferens. This 
procedure is called a vasectomy, and it is an effective form of male birth 
control. Although it may be possible to reverse a vasectomy, clinicians 


consider the procedure permanent, and advise men to undergo it only if they 
are certain they no longer wish to father children. 


Note: 
Interactive Link Feature 


eas 


— openstax COLLEGE” 


aun 


Watch this video to learn about a vasectomy. As described in this video, a 
vasectomy is a procedure in which a small section of the ductus (vas) 
deferens is removed from the scrotum. This interrupts the path taken by 
sperm through the ductus deferens. If sperm do not exit through the vas, 
either because the man has had a vasectomy or has not ejaculated, in what 
region of the testis do they remain? 


From each epididymis, each ductus deferens extends superiorly into the 
abdominal cavity through the inguinal canal in the abdominal wall. From 
here, the ductus deferens continues posteriorly to the pelvic cavity, ending 
posterior to the bladder where it dilates in a region called the ampulla 
(meaning “flask”). 


Sperm make up only 5 percent of the final volume of semen, the thick, 
milky fluid that the male ejaculates. The bulk of semen is produced by three 
critical accessory glands of the male reproductive system: the seminal 
vesicles, the prostate, and the bulbourethral glands. 


Seminal Vesicles 


As sperm pass through the ampulla of the ductus deferens at ejaculation, 
they mix with fluid from the associated seminal vesicle (see [link]). The 
paired seminal vesicles are glands that contribute approximately 60 percent 
of the semen volume. Seminal vesicle fluid contains large amounts of 
fructose, which is used by the sperm mitochondria to generate ATP to allow 
movement through the female reproductive tract. 


The fluid, now containing both sperm and seminal vesicle secretions, next 
moves into the associated ejaculatory duct, a short structure formed from 
the ampulla of the ductus deferens and the duct of the seminal vesicle. The 
paired ejaculatory ducts transport the seminal fluid into the next structure, 

the prostate gland. 


Prostate Gland 


As shown in [link], the centrally located prostate gland sits anterior to the 
rectum at the base of the bladder surrounding the prostatic urethra (the 
portion of the urethra that runs within the prostate). About the size of a 
walnut, the prostate is formed of both muscular and glandular tissues. It 
excretes an alkaline, milky fluid to the passing seminal fluid—now called 
semen—that is critical to first coagulate and then decoagulate the semen 
following ejaculation. The temporary thickening of semen helps retain it 
within the female reproductive tract, providing time for sperm to utilize the 
fructose provided by seminal vesicle secretions. When the semen regains its 
fluid state, sperm can then pass farther into the female reproductive tract. 


The prostate normally doubles in size during puberty. At approximately age 
25, it gradually begins to enlarge again. This enlargement does not usually 
cause problems; however, abnormal growth of the prostate, or benign 
prostatic hyperplasia (BPH), can cause constriction of the urethra as it 
passes through the middle of the prostate gland, leading to a number of 
lower urinary tract symptoms, such as a frequent and intense urge to 
urinate, a weak stream, and a sensation that the bladder has not emptied 
completely. By age 60, approximately 40 percent of men have some degree 
of BPH. By age 80, the number of affected individuals has jumped to as 
many as 80 percent. Treatments for BPH attempt to relieve the pressure on 


the urethra so that urine can flow more normally. Mild to moderate 
symptoms are treated with medication, whereas severe enlargement of the 
prostate is treated by surgery in which a portion of the prostate tissue is 
removed. 


Another common disorder involving the prostate is prostate cancer. 
According to the Centers for Disease Control and Prevention (CDC), 
prostate cancer is the second most common cancer in men. However, some 
forms of prostate cancer grow very slowly and thus may not ever require 
treatment. Aggressive forms of prostate cancer, in contrast, involve 
metastasis to vulnerable organs like the lungs and brain. There is no link 
between BPH and prostate cancer, but the symptoms are similar. Prostate 
cancer is detected by a medical history, a blood test, and a rectal exam that 
allows physicians to palpate the prostate and check for unusual masses. If a 
mass is detected, the cancer diagnosis is confirmed by biopsy of the cells. 


Bulbourethral Glands 


The final addition to semen is made by two bulbourethral glands (or 
Cowper’s glands) that release a thick, salty fluid that lubricates the end of 
the urethra and the vagina, and helps to clean urine residues from the penile 
urethra. The fluid from these accessory glands is released after the male 
becomes sexually aroused, and shortly before the release of the semen. It is 
therefore sometimes called pre-ejaculate. It is important to note that, in 
addition to the lubricating proteins, it is possible for bulbourethral fluid to 
pick up sperm already present in the urethra, and therefore it may be able to 
cause pregnancy. 


Note: 
Interactive Link Feature 


Watch this video to explore the structures of the male reproductive system 
and the path of sperm, which starts in the testes and ends as the sperm 
leave the penis through the urethra. Where are sperm deposited after they 
leave the ejaculatory duct? 


The Penis 


The penis is the male organ of copulation (sexual intercourse). It is flaccid 
for non-sexual actions, such as urination, and turgid and rod-like with 
sexual arousal. When erect, the stiffness of the organ allows it to penetrate 
into the vagina and deposit semen into the female reproductive tract. 
Cross-Sectional Anatomy of the Penis 


Flaccid: Lateral view Flaccid: Transverse view 


Penile venules 
(uncompressed) 


Deep dorsal vein 
Corpora cavernosa 
Cavernosal arteries 


Spongy urethra 


Prepuce 
Corpus spongiosum 


Erect: Transverse view 


Erect: Lateral view 


(4) Cavernosal arteries dilate, 
engorging corporal tissue 
with blood 


@ Engorging causes corporal tissue 
to swell, erecting the penis 


Engorged corporal tissue compresses penile 
veins and venules, maintaining erection 


Three columns of erectile tissue make up most of the volume 
of the penis. 


The shaft of the penis surrounds the urethra ([link]). The shaft is composed 
of three column-like chambers of erectile tissue that span the length of the 
shaft. Each of the two larger lateral chambers is called a corpus 
cavernosum (plural = corpora cavernosa). Together, these make up the bulk 
of the penis. The corpus spongiosum, which can be felt as a raised ridge on 
the erect penis, is a smaller chamber that surrounds the spongy, or penile, 
urethra. The end of the penis, called the glans penis, has a high 
concentration of nerve endings, resulting in very sensitive skin that 
influences the likelihood of ejaculation (see [link]). The skin from the shaft 


extends down over the glans and forms a collar called the prepuce (or 
foreskin). The foreskin also contains a dense concentration of nerve 
endings, and both lubricate and protect the sensitive skin of the glans penis. 
A surgical procedure called circumcision, often performed for religious or 
social reasons, removes the prepuce, typically within days of birth. 


Both sexual arousal and REM sleep (during which dreaming occurs) can 
induce an erection. Penile erections are the result of vasocongestion, or 
engorgement of the tissues because of more arterial blood flowing into the 
penis than is leaving in the veins. During sexual arousal, nitric oxide (NO) 
is released from nerve endings near blood vessels within the corpora 
cavernosa and spongiosum. Release of NO activates a signaling pathway 
that results in relaxation of the smooth muscles that surround the penile 
arteries, causing them to dilate. This dilation increases the amount of blood 
that can enter the penis and induces the endothelial cells in the penile 
arterial walls to also secrete NO and perpetuate the vasodilation. The rapid 
increase in blood volume fills the erectile chambers, and the increased 
pressure of the filled chambers compresses the thin-walled penile venules, 
preventing venous drainage of the penis. The result of this increased blood 
flow to the penis and reduced blood return from the penis is erection. 
Depending on the flaccid dimensions of a penis, it can increase in size 
slightly or greatly during erection, with the average length of an erect penis 
measuring approximately 15 cm. 


Note: 

Disorders of the... Feature 

Male Reproductive System 

Erectile dysfunction (ED) is a condition in which a man has difficulty 
either initiating or maintaining an erection. The combined prevalence of 
minimal, moderate, and complete ED is approximately 40 percent in men 
at age 40, and reaches nearly 70 percent by 70 years of age. In addition to 
aging, ED is associated with diabetes, vascular disease, psychiatric 
disorders, prostate disorders, the use of some drugs such as certain 
antidepressants, and problems with the testes resulting in low testosterone 
concentrations. These physical and emotional conditions can lead to 


interruptions in the vasodilation pathway and result in an inability to 
achieve an erection. 

Recall that the release of NO induces relaxation of the smooth muscles that 
surround the penile arteries, leading to the vasodilation necessary to 
achieve an erection. To reverse the process of vasodilation, an enzyme 
called phosphodiesterase (PDE) degrades a key component of the NO 
signaling pathway called cGMP. There are several different forms of this 
enzyme, and PDE type 5 is the type of PDE found in the tissues of the 
penis. Scientists discovered that inhibiting PDE5 increases blood flow, and 
allows vasodilation of the penis to occur. 

PDEs and the vasodilation signaling pathway are found in the vasculature 
in other parts of the body. In the 1990s, clinical trials of a PDES inhibitor 
called sildenafil were initiated to treat hypertension and angina pectoris 
(chest pain caused by poor blood flow through the heart). The trial showed 
that the drug was not effective at treating heart conditions, but many men 
experienced erection and priapism (erection lasting longer than 4 hours). 
Because of this, a clinical trial was started to investigate the ability of 
sildenafil to promote erections in men suffering from ED. In 1998, the 
FDA approved the drug, marketed as Viagra®. Since approval of the drug, 
sildenafil and similar PDE inhibitors now generate over a billion dollars a 
year in sales, and are reported to be effective in treating approximately 70 
to 85 percent of cases of ED. Importantly, men with health problems— 
especially those with cardiac disease taking nitrates—should avoid Viagra 
or talk to their physician to find out if they are a candidate for the use of 
this drug, as deaths have been reported for at-risk users. 


Testosterone 


Testosterone, an androgen, is a steroid hormone produced by Leydig cells. 
The alternate term for Leydig cells, interstitial cells, reflects their location 
between the seminiferous tubules in the testes. In male embryos, 
testosterone is secreted by Leydig cells by the seventh week of 
development, with peak concentrations reached in the second trimester. 
This early release of testosterone results in the anatomical differentiation of 
the male sexual organs. In childhood, testosterone concentrations are low. 


They increase during puberty, activating characteristic physical changes and 
initiating spermatogenesis. 


Functions of Testosterone 


The continued presence of testosterone is necessary to keep the male 
reproductive system working properly, and Leydig cells produce 
approximately 6 to 7 mg of testosterone per day. Testicular steroidogenesis 
(the manufacture of androgens, including testosterone) results in 
testosterone concentrations that are 100 times higher in the testes than in the 
circulation. Maintaining these normal concentrations of testosterone 
promotes spermatogenesis, whereas low levels of testosterone can lead to 
infertility. In addition to intratesticular secretion, testosterone is also 
released into the systemic circulation and plays an important role in muscle 
development, bone growth, the development of secondary sex 
characteristics, and maintaining libido (sex drive) in both males and 
females. In females, the ovaries secrete small amounts of testosterone, 
although most is converted to estradiol. A small amount of testosterone is 
also secreted by the adrenal glands in both sexes. 


Control of Testosterone 


The regulation of testosterone concentrations throughout the body is critical 
for male reproductive function. The intricate interplay between the 
endocrine system and the reproductive system is shown in [link]. 
Regulation of Testosterone Production 


(1) Hypothalamus releases GnRH. 
GnRH stimulates the anterior pituitary 
to release FSH and LH. 


(8) inhibin negatively 
feeds back to anterior 
pituitary, inhibiting 
further release of FSH. 
Testosterone negatively 
feeds back to the 
hypothalamus and 
pituitary, inhibiting 
further release of 
GnRH, FSH, and LH. 


FSH release 
LH release 
@) LH stimulates 


the Leydig cells to 
release testosterone. 


Leydig (interstitial) cells 
— FSH stimulates the 

3 Sertoli cells to release 
ABP. ABP binds to 
testosterone, keeping 
the latter at a high 
concentration. 


Ne rails 


Inhibin release 


JOD) 

“f= ~ = Seminiferous 
— tubule 
Sertoli cell 


Androgen-binding Testosterone 
protein (ABP) release release 


The hypothalamus and pituitary gland regulate the production 
of testosterone and the cells that assist in spermatogenesis. 
GnRH activates the anterior pituitary to produce LH and FSH, 
which in turn stimulate Leydig cells and Sertoli cells, 
respectively. The system is a negative feedback loop because 
the end products of the pathway, testosterone and inhibin, 
interact with the activity of GnRH to inhibit their own 
production. 


The regulation of Leydig cell production of testosterone begins outside of 
the testes. The hypothalamus and the pituitary gland in the brain integrate 
external and internal signals to control testosterone synthesis and secretion. 
The regulation begins in the hypothalamus. Pulsatile release of a hormone 
called gonadotropin-releasing hormone (GnRH) from the hypothalamus 
stimulates the endocrine release of hormones from the pituitary gland. 
Binding of GnRH to its receptors on the anterior pituitary gland stimulates 
release of the two gonadotropins: luteinizing hormone (LH) and follicle- 
stimulating hormone (FSH). These two hormones are critical for 


reproductive function in both men and women. In men, FSH binds 
predominantly to the Sertoli cells within the seminiferous tubules to 
promote spermatogenesis. FSH also stimulates the Sertoli cells to produce 
hormones called inhibins, which function to inhibit FSH release from the 
pituitary, thus reducing testosterone secretion. These polypeptide hormones 
correlate directly with Sertoli cell function and sperm number; inhibin B 
can be used as a marker of spermatogenic activity. In men, LH binds to 
receptors on Leydig cells in the testes and upregulates the production of 
testosterone. 


A negative feedback loop predominantly controls the synthesis and 
secretion of both FSH and LH. Low blood concentrations of testosterone 
stimulate the hypothalamic release of GnRH. GnRH then stimulates the 
anterior pituitary to secrete LH into the bloodstream. In the testis, LH binds 
to LH receptors on Leydig cells and stimulates the release of testosterone. 
When concentrations of testosterone in the blood reach a critical threshold, 
testosterone itself will bind to androgen receptors on both the hypothalamus 
and the anterior pituitary, inhibiting the synthesis and secretion of GnRH 
and LH, respectively. When the blood concentrations of testosterone once 
again decline, testosterone no longer interacts with the receptors to the same 
degree and GnRH and LH are once again secreted, stimulating more 
testosterone production. This same process occurs with FSH and inhibin to 
control spermatogenesis. 


Note: 

Aging and the... Feature 

Male Reproductive System 

Declines in Leydig cell activity can occur in men beginning at 40 to 50 
years of age. The resulting reduction in circulating testosterone 
concentrations can lead to symptoms of andropause, also known as male 
menopause. While the reduction in sex steroids in men is akin to female 
menopause, there is no clear sign—such as a lack of a menstrual period— 
to denote the initiation of andropause. Instead, men report feelings of 
fatigue, reduced muscle mass, depression, anxiety, irritability, loss of 
libido, and insomnia. A reduction in spermatogenesis resulting in lowered 


fertility is also reported, and sexual dysfunction can also be associated with 
andropausal symptoms. 

Whereas some researchers believe that certain aspects of andropause are 
difficult to tease apart from aging in general, testosterone replacement is 
sometimes prescribed to alleviate some symptoms. Recent studies have 
shown a benefit from androgen replacement therapy on the new onset of 
depression in elderly men; however, other studies caution against 
testosterone replacement for long-term treatment of andropause symptoms, 
showing that high doses can sharply increase the risk of both heart disease 
and prostate cancer. 


Chapter Review 


Gametes are the reproductive cells that combine to form offspring. Organs 
called gonads produce the gametes, along with the hormones that regulate 
human reproduction. The male gametes are called sperm. Spermatogenesis, 
the production of sperm, occurs within the seminiferous tubules that make 
up most of the testis. The scrotum is the muscular sac that holds the testes 
outside of the body cavity. 


Spermatogenesis begins with mitotic division of spermatogonia (stem cells) 
to produce primary spermatocytes that undergo the two divisions of meiosis 
to become secondary spermatocytes, then the haploid spermatids. During 
spermiogenesis, spermatids are transformed into spermatozoa (formed 
sperm). Upon release from the seminiferous tubules, sperm are moved to 
the epididymis where they continue to mature. During ejaculation, sperm 
exit the epididymis through the ductus deferens, a duct in the spermatic 
cord that leaves the scrotum. The ampulla of the ductus deferens meets the 
seminal vesicle, a gland that contributes fructose and proteins, at the 
ejaculatory duct. The fluid continues through the prostatic urethra, where 
secretions from the prostate are added to form semen. These secretions help 
the sperm to travel through the urethra and into the female reproductive 
tract. Secretions from the bulbourethral glands protect sperm and cleanse 
and lubricate the penile (spongy) urethra. 


The penis is the male organ of copulation. Columns of erectile tissue called 
the corpora cavernosa and corpus spongiosum fill with blood when sexual 
arousal activates vasodilatation in the blood vessels of the penis. 
Testosterone regulates and maintains the sex organs and sex drive, and 
induces the physical changes of puberty. Interplay between the testes and 
the endocrine system precisely control the production of testosterone with a 
negative feedback loop. 


Interactive Link Questions 


Exercise: 


Problem: 


Watch this video to learn about vasectomy. As described in this video, 
a vasectomy is a procedure in which a small section of the ductus (vas) 
deferens is removed from the scrotum. This interrupts the path taken 
by sperm through the ductus deferens. If sperm do not exit through the 
vas, either because the man has had a vasectomy or has not ejaculated, 
in what region of the testis do they remain? 


Solution: 


Sperm remain in the epididymis until they degenerate. 
Exercise: 
Problem: 
Watch this video to explore the structures of the male reproductive 
system and the path of sperm that starts in the testes and ends as the 


sperm leave the penis through the urethra. Where are sperm deposited 
after they leave the ejaculatory duct? 


Solution: 


Sperm enter the prostate. 


Review Questions 


Exercise: 


Problem: What are male gametes called? 


a. OVa 
b. sperm 
c. testes 
d. testosterone 


Solution: 


b 


Exercise: 


Problem: Leydig cells 


a. secrete testosterone 

b. activate the sperm flagellum 
c. support spermatogenesis 

d. secrete seminal fluid 


Solution: 


A 
Exercise: 
Problem: 


Which hypothalamic hormone contributes to the regulation of the male 
reproductive system? 


a. luteinizing hormone 
b. gonadotropin-releasing hormone 


c. follicle-stimulating hormone 
d. androgens 

Solution: 

b 

Exercise: 

Problem: What is the function of the epididymis? 
a. sperm maturation and storage 
b. produces the bulk of seminal fluid 


c. provides nitric oxide needed for erections 
d. spermatogenesis 


Solution: 
a 
Exercise: 
Problem: Spermatogenesis takes place in the 
a. prostate gland 
b. glans penis 


c. seminiferous tubules 
d. ejaculatory duct 


Solution: 


Critical Thinking Questions 


Exercise: 
Problem: 


Briefly explain why mature gametes carry only one set of 
chromosomes. 


Solution: 


A single gamete must combine with a gamete from an individual of the 
opposite sex to produce a fertilized egg, which has a complete set of 
chromosomes and is the first cell of a new individual. 


Exercise: 
Problem: 


What special features are evident in sperm cells but not in somatic 
cells, and how do these specializations function? 


Solution: 


Unlike somatic cells, sperm are haploid. They also have very little 
cytoplasm. They have a head with a compact nucleus covered by an 
acrosome filled with enzymes, and a mid-piece filled with 
mitochondria that power their movement. They are motile because of 
their tail, a structure containing a flagellum, which is specialized for 
movement. 


Exercise: 
Problem: 


What do each of the three male accessory glands contribute to the 
semen? 


Solution: 
The three accessory glands make the following contributions to semen: 


the seminal vesicle contributes about 60 percent of the semen volume, 
with fluid that contains large amounts of fructose to power the 


movement of sperm; the prostate gland contributes substances critical 
to sperm maturation; and the bulbourethral glands contribute a thick 
fluid that lubricates the ends of the urethra and the vagina and helps to 
clean urine residues from the urethra. 


Exercise: 


Problem: Describe how penile erection occurs. 


Solution: 


During sexual arousal, nitric oxide (NO) is released from nerve 
endings near blood vessels within the corpora cavernosa and corpus 
spongiosum. The release of NO activates a signaling pathway that 
results in relaxation of the smooth muscles that surround the penile 
arteries, causing them to dilate. This dilation increases the amount of 
blood that can enter the penis, and induces the endothelial cells in the 
penile arterial walls to secrete NO, perpetuating the vasodilation. The 
rapid increase in blood volume fills the erectile chambers, and the 
increased pressure of the filled chambers compresses the thin-walled 
penile venules, preventing venous drainage of the penis. An erection is 
the result of this increased blood flow to the penis and reduced blood 
return from the penis. 


Exercise: 


Problem: 


While anabolic steroids (synthetic testosterone) bulk up muscles, they 
can also affect testosterone production in the testis. Using what you 
know about negative feedback, describe what would happen to 
testosterone production in the testis if a male takes large amounts of 
synthetic testosterone. 


Solution: 


Testosterone production by the body would be reduced if a male were 
taking anabolic steroids. This is because the hypothalamus responds to 
rising testosterone levels by reducing its secretion of GnRH, which 


would in turn reduce the anterior pituitary’s release of LH, finally 
reducing the manufacture of testosterone in the testes. 


Glossary 


blood-testis barrier 
tight junctions between Sertoli cells that prevent bloodborne pathogens 
from gaining access to later stages of spermatogenesis and prevent the 
potential for an autoimmune reaction to haploid sperm 


bulbourethral glands 
(also, Cowper’s glands) glands that secrete a lubricating mucus that 
cleans and lubricates the urethra prior to and during ejaculation 


corpus cavernosum 
either of two columns of erectile tissue in the penis that fill with blood 
during an erection 


corpus spongiosum 
(plural = corpora cavernosa) column of erectile tissue in the penis that 
fills with blood during an erection and surrounds the penile urethra on 
the ventral portion of the penis 


ductus deferens 
(also, vas deferens) duct that transports sperm from the epididymis 
through the spermatic cord and into the ejaculatory duct; also referred 
as the vas deferens 


ejaculatory duct 
duct that connects the ampulla of the ductus deferens with the duct of 
the seminal vesicle at the prostatic urethra 


epididymis 
(plural = epididymides) coiled tubular structure in which sperm start to 


mature and are stored until ejaculation 


gamete 


haploid reproductive cell that contributes genetic material to form an 
offspring 


glans penis 
bulbous end of the penis that contains a large number of nerve endings 


gonadotropin-releasing hormone (GnRH) 
hormone released by the hypothalamus that regulates the production of 
follicle-stimulating hormone and luteinizing hormone from the 
pituitary gland 


gonads 
reproductive organs (testes in men and ovaries in women) that produce 
gametes and reproductive hormones 


inguinal canal 
opening in abdominal wall that connects the testes to the abdominal 
cavity 


Leydig cells 
cells between the seminiferous tubules of the testes that produce 
testosterone; a type of interstitial cell 


penis 
male organ of copulation 


prepuce 
(also, foreskin) flap of skin that forms a collar around, and thus 
protects and lubricates, the glans penis; also referred as the foreskin 


prostate gland 
doughnut-shaped gland at the base of the bladder surrounding the 
urethra and contributing fluid to semen during ejaculation 


scrotum 
external pouch of skin and muscle that houses the testes 


semen 


ejaculatory fluid composed of sperm and secretions from the seminal 
vesicles, prostate, and bulbourethral glands 


seminal vesicle 
gland that produces seminal fluid, which contributes to semen 


seminiferous tubules 
tube structures within the testes where spermatogenesis occurs 


Sertoli cells 
cells that support germ cells through the process of spermatogenesis; a 
type of sustentacular cell 


sperm 
(also, spermatozoon) male gamete 


spermatic cord 
bundle of nerves and blood vessels that supplies the testes; contains 
ductus deferens 


spermatid 
immature sperm cells produced by meiosis II of secondary 
spermatocytes 


spermatocyte 
cell that results from the division of spermatogonium and undergoes 
meiosis I and meiosis IT to form spermatids 


spermatogenesis 
formation of new sperm, occurs in the seminiferous tubules of the 
testes 


spermatogonia 
(singular = spermatogonium) diploid precursor cells that become 
sperm 


spermiogenesis 
transformation of spermatids to spermatozoa during spermatogenesis 


testes 
(singular = testis) male gonads 


Female Anatomy 
By the end of this section, you will be able to: 


¢ Describe the structure and function of the organs of the female 
reproductive system 

e List the steps of oogenesis 

e Describe the hormonal changes that occur during the ovarian and 
menstrual cycles 

e Trace the path of an oocyte from ovary to fertilization 


The female reproductive system functions to produce gametes and 
reproductive hormones, just like the male reproductive system; however, it 
also has the additional task of supporting the developing fetus and 
delivering it to the outside world. Unlike its male counterpart, the female 
reproductive system is located primarily inside the pelvic cavity ([link]). 
Recall that the ovaries are the female gonads. The gamete they produce is 
called an oocyte. We’|I discuss the production of oocytes in detail shortly. 
First, let’s look at some of the structures of the female reproductive system. 
Female Reproductive System 


Uterus 
Ovary 


Bladder 
Pubic sym is Fornix of uterus 
Mons pubis Cervix 
Urethra Rectum 
Clitoris Vagina 
Labium minora 
Anus 
Labium majora 
(a) Human female reproductive system: lateral view 
Ovary 
Fimbriae 
Ovarian 
earns ligament 
=s_4 5 Broad 
= = SS ‘ ligament 
(oviduct) 
Cervix 
Vagina 


(b) Human female reproductive system: anterior view 


The major organs of the female 
reproductive system are located inside the 
pelvic cavity. 


External Female Genitals 


The external female reproductive structures are referred to collectively as 
the vulva ({link]). The mons pubis is a pad of fat that is located at the 
anterior, over the pubic bone. After puberty, it becomes covered in pubic 
hair. The labia majora (labia = “lips”; majora = “larger”) are folds of hair- 


covered skin that begin just posterior to the mons pubis. The thinner and 
more pigmented labia minora (labia = “lips”; minora = “smaller”) extend 
medial to the labia majora. Although they naturally vary in shape and size 
from woman to woman, the labia minora serve to protect the female urethra 
and the entrance to the female reproductive tract. 


The superior, anterior portions of the labia minora come together to encircle 
the clitoris (or glans clitoris), an organ that originates from the same cells 
as the glans penis and has abundant nerves that make it important in sexual 
sensation and orgasm. The hymen is a thin membrane that sometimes 
partially covers the entrance to the vagina. An intact hymen cannot be used 
as an indication of “virginity”; even at birth, this is only a partial 
membrane, as menstrual fluid and other secretions must be able to exit the 
body, regardless of penile—vaginal intercourse. The vaginal opening is 
located between the opening of the urethra and the anus. It is flanked by 
outlets to the Bartholin’s glands (or greater vestibular glands). 

The Vulva 


Corpus cavernosum 


J 
ot 

—____" 

- Urethral opening — + hs 

= Labia majora ii ‘a 

: 2 } 7 

FL ‘ 

3 ' 


Bulb of vestibule 


Opening of right Ba . y ee 


Bartholin's gland 


/ 


A= Anus 
AN Bartholin’s glands 
Vulva: External anterior view Vulva: Internal anteriolateral view 


The external female genitalia are referred to collectively as the 
vulva. 


Vagina 


The vagina, shown at the bottom of [link] and [link], is a muscular canal 
(approximately 10 cm long) that serves as the entrance to the reproductive 
tract. It also serves as the exit from the uterus during menses and childbirth. 
The outer walls of the anterior and posterior vagina are formed into 
longitudinal columns, or ridges, and the superior portion of the vagina— 
called the fornix—meets the protruding uterine cervix. The walls of the 
vagina are lined with an outer, fibrous adventitia; a middle layer of smooth 
muscle; and an inner mucous membrane with transverse folds called rugae. 
Together, the middle and inner layers allow the expansion of the vagina to 
accommodate intercourse and childbirth. The thin, perforated hymen can 
partially surround the opening to the vaginal orifice. The hymen can be 
ruptured with strenuous physical exercise, penile—vaginal intercourse, and 
childbirth. The Bartholin’s glands and the lesser vestibular glands (located 
near the clitoris) secrete mucus, which keeps the vestibular area moist. 


The vagina is home to a normal population of microorganisms that help to 
protect against infection by pathogenic bacteria, yeast, or other organisms 
that can enter the vagina. In a healthy woman, the most predominant type of 
vaginal bacteria is from the genus Lactobacillus. This family of beneficial 
bacterial flora secretes lactic acid, and thus protects the vagina by 
maintaining an acidic pH (below 4.5). Potential pathogens are less likely to 
survive in these acidic conditions. Lactic acid, in combination with other 
vaginal secretions, makes the vagina a self-cleansing organ. However, 
douching—or washing out the vagina with fluid—can disrupt the normal 
balance of healthy microorganisms, and actually increase a woman’s risk 
for infections and irritation. Indeed, the American College of Obstetricians 
and Gynecologists recommend that women do not douche, and that they 
allow the vagina to maintain its normal healthy population of protective 
microbial flora. 


Ovaries 


The ovaries are the female gonads (see [link]). Paired ovals, they are each 
about 2 to 3 cm in length, about the size of an almond. The ovaries are 
located within the pelvic cavity, and are supported by the mesovarium, an 


extension of the peritoneum that connects the ovaries to the broad 
ligament. Extending from the mesovarium itself is the suspensory ligament 
that contains the ovarian blood and lymph vessels. Finally, the ovary itself 
is attached to the uterus via the ovarian ligament. 


The ovary comprises an outer covering of cuboidal epithelium called the 
ovarian surface epithelium that is superficial to a dense connective tissue 
covering called the tunica albuginea. Beneath the tunica albuginea is the 
cortex, or outer portion, of the organ. The cortex is composed of a tissue 
framework called the ovarian stroma that forms the bulk of the adult ovary. 
Oocytes develop within the outer layer of this stroma, each surrounded by 
supporting cells. This grouping of an oocyte and its supporting cells is 
called a follicle. The growth and development of ovarian follicles will be 
described shortly. Beneath the cortex lies the inner ovarian medulla, the site 
of blood vessels, lymph vessels, and the nerves of the ovary. You will learn 
more about the overall anatomy of the female reproductive system at the 
end of this section. 


The Ovarian Cycle 


The ovarian cycle is a set of predictable changes in a female’s oocytes and 
ovarian follicles. During a woman’s reproductive years, it is a roughly 28- 
day cycle that can be correlated with, but is not the same as, the menstrual 
cycle (discussed shortly). The cycle includes two interrelated processes: 
oogenesis (the production of female gametes) and folliculogenesis (the 
growth and development of ovarian follicles). 


Oogenesis 


Gametogenesis in females is called oogenesis. The process begins with the 
Ovarian stem cells, or oogonia (({link]). Oogonia are formed during fetal 
development, and divide via mitosis, much like spermatogonia in the testis. 
Unlike spermatogonia, however, oogonia form primary oocytes in the fetal 
ovary prior to birth. These primary oocytes are then arrested in this stage of 
meiosis I, only to resume it years later, beginning at puberty and continuing 


until the woman is near menopause (the cessation of a woman’s 
reproductive functions). The number of primary oocytes present in the 
ovaries declines from one to two million in an infant, to approximately 
400,000 at puberty, to zero by the end of menopause. 


The initiation of ovulation—the release of an oocyte from the ovary— 
marks the transition from puberty into reproductive maturity for women. 
From then on, throughout a woman’s reproductive years, ovulation occurs 
approximately once every 28 days. Just prior to ovulation, a surge of 
luteinizing hormone triggers the resumption of meiosis in a primary oocyte. 
This initiates the transition from primary to secondary oocyte. However, as 
you can see in [link], this cell division does not result in two identical cells. 
Instead, the cytoplasm is divided unequally, and one daughter cell is much 
larger than the other. This larger cell, the secondary oocyte, eventually 
leaves the ovary during ovulation. The smaller cell, called the first polar 
body, may or may not complete meiosis and produce second polar bodies; 
in either case, it eventually disintegrates. Therefore, even though oogenesis 
produces up to four cells, only one survives. 

Oogenesis 


Te. 
— (*) oO By onto 


Meiosis arrests in 
t prophase | 
Before birth 


After pube! 
in m | 3 Meiosis | resumes 


Oocyte meiosis 
arrests at Secondary First polar 
metaphase II oocyte body 


Before sperm penetration 


After sperm penetration 
4 


Oocyte meiosis Second polar 
completes bodies 
immediately after 

sperm penetrates 

the oocyte 


The unequal cell division of oogenesis produces one to three 
polar bodies that later degrade, as well as a single haploid 
ovum, which is produced only if there is penetration of the 

secondary oocyte by a sperm cell. 


How does the diploid secondary oocyte become an ovum—the haploid 
female gamete? Meiosis of a secondary oocyte is completed only if a sperm 
succeeds in penetrating its barriers. Meiosis II then resumes, producing one 
haploid ovum that, at the instant of fertilization by a (haploid) sperm, 
becomes the first diploid cell of the new offspring (a zygote). Thus, the 
ovum can be thought of as a brief, transitional, haploid stage between the 
diploid oocyte and diploid zygote. 


The larger amount of cytoplasm contained in the female gamete is used to 
supply the developing zygote with nutrients during the period between 
fertilization and implantation into the uterus. Interestingly, sperm contribute 
only DNA at fertilization —not cytoplasm. Therefore, the cytoplasm and all 
of the cytoplasmic organelles in the developing embryo are of maternal 
origin. This includes mitochondria, which contain their own DNA. 
Scientific research in the 1980s determined that mitochondrial DNA was 
maternally inherited, meaning that you can trace your mitochondrial DNA 
directly to your mother, her mother, and so on back through your female 
ancestors. 


Note: 

Everyday Connections Feature 

Mapping Human History with Mitochondrial DNA 

When we talk about human DNA, we’re usually referring to nuclear DNA; 
that is, the DNA coiled into chromosomal bundles in the nucleus of our 
cells. We inherit half of our nuclear DNA from our father, and half from 
our mother. However, mitochondrial DNA (mtDNA) comes only from the 
mitochondria in the cytoplasm of the fat ovum we inherit from our mother. 
She received her mtDNA from her mother, who got it from her mother, and 
so on. Each of our cells contains approximately 1700 mitochondria, with 
each mitochondrion packed with mtDNA containing approximately 37 
genes. 

Mutations (changes) in mtDNA occur spontaneously in a somewhat 
organized pattern at regular intervals in human history. By analyzing these 
mutational relationships, researchers have been able to determine that we 
can all trace our ancestry back to one woman who lived in Africa about 
200,000 years ago. Scientists have given this woman the biblical name 
Eve, although she is not, of course, the first Homo sapiens female. More 
precisely, she is our most recent common ancestor through matrilineal 
descent. 

This doesn’t mean that everyone’s mtDNA today looks exactly like that of 
our ancestral Eve. Because of the spontaneous mutations in mtDNA that 
have occurred over the centuries, researchers can map different “branches” 
off of the “main trunk” of our mtDNA family tree. Your mtDNA might 


have a pattern of mutations that aligns more closely with one branch, and 
your neighbor’s may align with another branch. Still, all branches 
eventually lead back to Eve. 

But what happened to the mtDNA of all of the other Homo sapiens females 
who were living at the time of Eve? Researchers explain that, over the 
centuries, their female descendants died childless or with only male 
children, and thus, their maternal line—and its mtDNA—ended. 


Folliculogenesis 


Again, ovarian follicles are oocytes and their supporting cells. They grow 
and develop in a process called folliculogenesis, which typically leads to 
ovulation of one follicle approximately every 28 days, along with death to 
multiple other follicles. The death of ovarian follicles is called atresia, and 
can occur at any point during follicular development. Recall that, a female 
infant at birth will have one to two million oocytes within her ovarian 
follicles, and that this number declines throughout life until menopause, 
when no follicles remain. As you’ll see next, follicles progress from 
primordial, to primary, to secondary and tertiary stages prior to ovulation— 
with the oocyte inside the follicle remaining as a primary oocyte until right 
before ovulation. 


Folliculogenesis begins with follicles in a resting state. These small 
primordial follicles are present in newborn females and are the prevailing 
follicle type in the adult ovary ([link]). Primordial follicles have only a 
single flat layer of support cells, called granulosa cells, that surround the 
oocyte, and they can stay in this resting state for years—some until right 
before menopause. 


After puberty, a few primordial follicles will respond to a recruitment signal 
each day, and will join a pool of immature growing follicles called primary 
follicles. Primary follicles start with a single layer of granulosa cells, but 
the granulosa cells then become active and transition from a flat or 
Squamous shape to a rounded, cuboidal shape as they increase in size and 
proliferate. As the granulosa cells divide, the follicles—now called 
secondary follicles (see [link ])—increase in diameter, adding a new outer 


layer of connective tissue, blood vessels, and theca cells—cells that work 
with the granulosa cells to produce estrogens. 


Within the growing secondary follicle, the primary oocyte now secretes a 
thin acellular membrane called the zona pellucida that will play a critical 
role in fertilization. A thick fluid, called follicular fluid, that has formed 
between the granulosa cells also begins to collect into one large pool, or 
antrum. Follicles in which the antrum has become large and fully formed 
are considered tertiary follicles (or antral follicles). Several follicles reach 
the tertiary stage at the same time, and most of these will undergo atresia. 
The one that does not die will continue to grow and develop until ovulation, 
when it will expel its secondary oocyte surrounded by several layers of 
granulosa cells from the ovary. Keep in mind that most follicles don’t make 
it to this point. In fact, roughly 99 percent of the follicles in the ovary will 
undergo atresia, which can occur at any stage of folliculogenesis. 
Folliculogenesis 


(a) Stages of Folliculogenesis 
(2) Primordial follicle (@) Primary follicie (8) Secondary follicle 


Granulosa cells Oocyte Granulosa cells 


“| 
‘G) Corpus luteum 6) Ovulating follicle @) Tertiary follicle 
pi Granulosa cells 


roc 


(b) A Secondary Follicle 


(a) The maturation of a follicle is shown in a clockwise 
direction proceeding from the primordial follicles. FSH 
stimulates the growth of a tertiary follicle, and LH stimulates 
the production of estrogen by granulosa and theca cells. Once 
the follicle is mature, it ruptures and releases the oocyte. Cells 


remaining in the follicle then develop into the corpus luteum. 
(b) In this electron micrograph of a secondary follicle, the 
oocyte, theca cells (thecae folliculi), and developing antrum are 
clearly visible. EM x 1100. (Micrograph provided by the 
Regents of University of Michigan Medical School © 2012) 


Hormonal Control of the Ovarian Cycle 


The process of development that we have just described, from primordial 
follicle to early tertiary follicle, takes approximately two months in humans. 
The final stages of development of a small cohort of tertiary follicles, 
ending with ovulation of a secondary oocyte, occur over a course of 
approximately 28 days. These changes are regulated by many of the same 
hormones that regulate the male reproductive system, including GnRH, LH, 
and FSH. 


As in men, the hypothalamus produces GnRH, a hormone that signals the 
anterior pituitary gland to produce the gonadotropins FSH and LH ([lLink]). 
These gonadotropins leave the pituitary and travel through the bloodstream 
to the ovaries, where they bind to receptors on the granulosa and theca cells 
of the follicles. FSH stimulates the follicles to grow (hence its name of 
follicle-stimulating hormone), and the five or six tertiary follicles expand in 
diameter. The release of LH also stimulates the granulosa and theca cells of 
the follicles to produce the sex steroid hormone estradiol, a type of 
estrogen. This phase of the ovarian cycle, when the tertiary follicles are 
growing and secreting estrogen, is known as the follicular phase. 


The more granulosa and theca cells a follicle has (that is, the larger and 
more developed it is), the more estrogen it will produce in response to LH 
stimulation. As a result of these large follicles producing large amounts of 
estrogen, systemic plasma estrogen concentrations increase. Following a 
classic negative feedback loop, the high concentrations of estrogen will 
stimulate the hypothalamus and pituitary to reduce the production of GnRH, 
LH, and FSH. Because the large tertiary follicles require FSH to grow and 


survive at this point, this decline in FSH caused by negative feedback leads 
most of them to die (atresia). Typically only one follicle, now called the 
dominant follicle, will survive this reduction in FSH, and this follicle will 
be the one that releases an oocyte. Scientists have studied many factors that 
lead to a particular follicle becoming dominant: size, the number of 
granulosa cells, and the number of FSH receptors on those granulosa cells 
all contribute to a follicle becoming the one surviving dominant follicle. 
Hormonal Regulation of Ovulation 


@) Follicular phase @ ovulation 

Pituitary hormone Pituitary hormone 

effect: LH and FSH effect: LH and FSH 

stimulate several —” stimulate maturation cia 
follicles to grow. of one of the 


growing follicles. 


Estradiol 


FSH 
LH Estradio| Ovarian a Ovarian 
hormone hormone 
effects: effects: 
Boninat ll crowing lo 
produces estradiol, continues to produce 


Estradio! Which: 


estradiol, which: 
* Inhibits GnRH, FSH, Estradiol ". stimulates GnRH, 
and LH production FSH, and LH 
* Causes production 
endometrium to Endometrium + LH surge triggers 
thicken ovulation 


@)Luteal phase 


Pituitary hormone 
effect: LH stimulates 
formation of a corpus 
luteum from follicular 
tissue left behind after 
ovulation. 


| GnRH 


FSH 
LH 


Progesterone Ovarlan 
hormone 
ee 


The corpus 
luteum secretes 
progesterone, which: 
¢ Inhibits GnRH, FSH, 
Progesterone = and LH production 
ae * Maintains the 
aaa 
corpus luteum degrades, 
progesterone declines, 
initiating sloughing of 
the stratum functionalis 


The hypothalamus and pituitary gland regulate the ovarian 
cycle and ovulation. GnRH activates the anterior pituitary to 
produce LH and FSH, which stimulate the production of 
estrogen and progesterone by the ovaries. 


When only the one dominant follicle remains in the ovary, it again begins to 
secrete estrogen. It produces more estrogen than all of the developing 
follicles did together before the negative feedback occurred. It produces so 
much estrogen that the normal negative feedback doesn’t occur. Instead, 
these extremely high concentrations of systemic plasma estrogen trigger a 
regulatory switch in the anterior pituitary that responds by secreting large 
amounts of LH and FSH into the bloodstream (see [link]). The positive 
feedback loop by which more estrogen triggers release of more LH and 
FSH only occurs at this point in the cycle. 


It is this large burst of LH (called the LH surge) that leads to ovulation of 
the dominant follicle. The LH surge induces many changes in the dominant 
follicle, including stimulating the resumption of meiosis of the primary 
oocyte to a secondary oocyte. As noted earlier, the polar body that results 
from unequal cell division simply degrades. The LH surge also triggers 
proteases (enzymes that cleave proteins) to break down structural proteins 
in the ovary wall on the surface of the bulging dominant follicle. This 
degradation of the wall, combined with pressure from the large, fluid-filled 
antrum, results in the expulsion of the oocyte surrounded by granulosa cells 
into the peritoneal cavity. This release is ovulation. 


In the next section, you will follow the ovulated oocyte as it travels toward 
the uterus, but there is one more important event that occurs in the ovarian 
cycle. The surge of LH also stimulates a change in the granulosa and theca 
cells that remain in the follicle after the oocyte has been ovulated. This 
change is called luteinization (recall that the full name of LH is luteinizing 
hormone), and it transforms the collapsed follicle into a new endocrine 
structure called the corpus luteum, a term meaning “yellowish body” (see 
[link]). Instead of estrogen, the luteinized granulosa and theca cells of the 
corpus luteum begin to produce large amounts of the sex steroid hormone 


progesterone, a hormone that is critical for the establishment and 
maintenance of pregnancy. Progesterone triggers negative feedback at the 
hypothalamus and pituitary, which keeps GnRH, LH, and FSH secretions 
low, so no new dominant follicles develop at this time. 


The post-ovulatory phase of progesterone secretion is known as the luteal 
phase of the ovarian cycle. If pregnancy does not occur within 10 to 12 
days, the corpus luteum will stop secreting progesterone and degrade into 
the corpus albicans, a nonfunctional “whitish body” that will disintegrate 
in the ovary over a period of several months. During this time of reduced 
progesterone secretion, FSH and LH are once again stimulated, and the 
follicular phase begins again with a new cohort of early tertiary follicles 
beginning to grow and secrete estrogen. 


The Uterine Tubes 


The uterine tubes (also called fallopian tubes or oviducts) serve as the 
conduit of the oocyte from the ovary to the uterus ([link]). Each of the two 
uterine tubes is close to, but not directly connected to, the ovary and divided 
into sections. The isthmus is the narrow medial end of each uterine tube 
that is connected to the uterus. The wide distal infundibulum flares out 
with slender, finger-like projections called fimbriae. The middle region of 
the tube, called the ampulla, is where fertilization often occurs. The uterine 
tubes also have three layers: an outer serosa, a middle smooth muscle layer, 
and an inner mucosal layer. In addition to its mucus-secreting cells, the 
inner mucosa contains ciliated cells that beat in the direction of the uterus, 
producing a current that will be critical to move the oocyte. 


Following ovulation, the secondary oocyte surrounded by a few granulosa 
cells is released into the peritoneal cavity. The nearby uterine tube, either 
left or right, receives the oocyte. Unlike sperm, oocytes lack flagella, and 
therefore cannot move on their own. So how do they travel into the uterine 
tube and toward the uterus? High concentrations of estrogen that occur 
around the time of ovulation induce contractions of the smooth muscle 
along the length of the uterine tube. These contractions occur every 4 to 8 
seconds, and the result is a coordinated movement that sweeps the surface 
of the ovary and the pelvic cavity. Current flowing toward the uterus is 


generated by coordinated beating of the cilia that line the outside and lumen 
of the length of the uterine tube. These cilia beat more strongly in response 
to the high estrogen concentrations that occur around the time of ovulation. 
As aresult of these mechanisms, the oocyte—granulosa cell complex is 
pulled into the interior of the tube. Once inside, the muscular contractions 
and beating cilia move the oocyte slowly toward the uterus. When 
fertilization does occur, sperm typically meet the egg while it is still moving 
through the ampulla. 


Note: 
Interactive Link 


CHES 


— 
mess Openstax COLLEGE 


cere 


Watch this video to observe ovulation and its initiation in response to the 
release of FSH and LH from the pituitary gland. What specialized 
structures help guide the oocyte from the ovary into the uterine tube? 


If the oocyte is successfully fertilized, the resulting zygote will begin to 
divide into two cells, then four, and so on, as it makes its way through the 
uterine tube and into the uterus. There, it will implant and continue to grow. 
If the egg is not fertilized, it will simply degrade—either in the uterine tube 
or in the uterus, where it may be shed with the next menstrual period. 
Ovaries, Uterine Tubes, and Uterus 


Uterine tube (oviduct) 


Infundibulum Ampulla Isthmus Fundus Broad 
ligament 


Edge of 
follicle 
Fimbriae 
Ovarian 
cortex and vein 
Suspensory 
ligament 


Tunica albuginea 


and vein 
Vaginal artery 
Vagina 


This anterior view shows the relationship of the ovaries, uterine 
tubes (oviducts), and uterus. Sperm enter through the vagina, 
and fertilization of an ovulated oocyte usually occurs in the 
distal uterine tube. From left to right, LM x 400, LM x 20. 
(Micrographs provided by the Regents of University of 
Michigan Medical School © 2012) 


The open-ended structure of the uterine tubes can have significant health 
consequences if bacteria or other contagions enter through the vagina and 
move through the uterus, into the tubes, and then into the pelvic cavity. If 
this is left unchecked, a bacterial infection (sepsis) could quickly become 
life-threatening. The spread of an infection in this manner is of special 
concern when unskilled practitioners perform abortions in non-sterile 
conditions. Sepsis is also associated with sexually transmitted bacterial 
infections, especially gonorrhea and chlamydia. These increase a woman’s 
risk for pelvic inflammatory disease (PID), infection of the uterine tubes or 
other reproductive organs. Even when resolved, PID can leave scar tissue in 
the tubes, leading to infertility. 


Note: 
Interactive Link 


Watch this series of videos to look at the movement of the oocyte through 
the ovary. The cilia in the uterine tube promote movement of the oocyte. 
What would likely occur if the cilia were paralyzed at the time of 
ovulation? 


The Uterus and Cervix 


The uterus is the muscular organ that nourishes and supports the growing 
embryo (see [link]). Its average size is approximately 5 cm wide by 7 cm 
long (approximately 2 in by 3 in) when a female is not pregnant. It has three 
sections. The portion of the uterus superior to the opening of the uterine 
tubes is called the fundus. The middle section of the uterus is called the 
body of uterus (or corpus). The cervix is the narrow inferior portion of the 
uterus that projects into the vagina. The cervix produces mucus secretions 
that become thin and stringy under the influence of high systemic plasma 
estrogen concentrations, and these secretions can facilitate sperm movement 
through the reproductive tract. 


Several ligaments maintain the position of the uterus within the 
abdominopelvic cavity. The broad ligament is a fold of peritoneum that 
serves as a primary support for the uterus, extending laterally from both 
sides of the uterus and attaching it to the pelvic wall. The round ligament 
attaches to the uterus near the uterine tubes, and extends to the labia majora. 
Finally, the uterosacral ligament stabilizes the uterus posteriorly by its 
connection from the cervix to the pelvic wall. 


The wall of the uterus is made up of three layers. The most superficial layer 
is the serous membrane, or perimetrium, which consists of epithelial tissue 
that covers the exterior portion of the uterus. The middle layer, or 


myometrium, is a thick layer of smooth muscle responsible for uterine 
contractions. Most of the uterus is myometrial tissue, and the muscle fibers 
run horizontally, vertically, and diagonally, allowing the powerful 
contractions that occur during labor and the less powerful contractions (or 
cramps) that help to expel menstrual blood during a woman’s period. 
Anteriorly directed myometrial contractions also occur near the time of 
ovulation, and are thought to possibly facilitate the transport of sperm 
through the female reproductive tract. 


The innermost layer of the uterus is called the endometrium. The 
endometrium contains a connective tissue lining, the lamina propria, which 
is covered by epithelial tissue that lines the lumen. Structurally, the 
endometrium consists of two layers: the stratum basalis and the stratum 
functionalis (the basal and functional layers). The stratum basalis layer is 
part of the lamina propria and is adjacent to the myometrium; this layer 
does not shed during menses. In contrast, the thicker stratum functionalis 
layer contains the glandular portion of the lamina propria and the 
endothelial tissue that lines the uterine lumen. It is the stratum functionalis 
that grows and thickens in response to increased levels of estrogen and 
progesterone. In the luteal phase of the menstrual cycle, special branches 
off of the uterine artery called spiral arteries supply the thickened stratum 
functionalis. This inner functional layer provides the proper site of 
implantation for the fertilized egg, and—should fertilization not occur—it is 
only the stratum functionalis layer of the endometrium that sheds during 
menstruation. 


Recall that during the follicular phase of the ovarian cycle, the tertiary 
follicles are growing and secreting estrogen. At the same time, the stratum 
functionalis of the endometrium is thickening to prepare for a potential 
implantation. The post-ovulatory increase in progesterone, which 
characterizes the luteal phase, is key for maintaining a thick stratum 
functionalis. As long as a functional corpus luteum is present in the ovary, 
the endometrial lining is prepared for implantation. Indeed, if an embryo 
implants, signals are sent to the corpus luteum to continue secreting 
progesterone to maintain the endometrium, and thus maintain the 
pregnancy. If an embryo does not implant, no signal is sent to the corpus 
luteum and it degrades, ceasing progesterone production and ending the 


luteal phase. Without progesterone, the endometrium thins and, under the 
influence of prostaglandins, the spiral arteries of the endometrium constrict 
and rupture, preventing oxygenated blood from reaching the endometrial 
tissue. As a result, endometrial tissue dies and blood, pieces of the 
endometrial tissue, and white blood cells are shed through the vagina during 
menstruation, or the menses. The first menses after puberty, called 
menarche, can occur either before or after the first ovulation. 


The Menstrual Cycle 


Now that we have discussed the maturation of the cohort of tertiary follicles 
in the ovary, the build-up and then shedding of the endometrial lining in the 
uterus, and the function of the uterine tubes and vagina, we can put 
everything together to talk about the three phases of the menstrual cycle— 
the series of changes in which the uterine lining is shed, rebuilds, and 
prepares for implantation. 


The timing of the menstrual cycle starts with the first day of menses, 
referred to as day one of a woman’s period. Cycle length is determined by 
counting the days between the onset of bleeding in two subsequent cycles. 
Because the average length of a woman’s menstrual cycle is 28 days, this is 
the time period used to identify the timing of events in the cycle. However, 
the length of the menstrual cycle varies among women, and even in the 
Same woman from one cycle to the next, typically from 21 to 32 days. 


Just as the hormones produced by the granulosa and theca cells of the ovary 
“drive” the follicular and luteal phases of the ovarian cycle, they also 
control the three distinct phases of the menstrual cycle. These are the 
menses phase, the proliferative phase, and the secretory phase. 


Menses Phase 


The menses phase of the menstrual cycle is the phase during which the 
lining is shed; that is, the days that the woman menstruates. Although it 
averages approximately five days, the menses phase can last from 2 to 7 
days, or longer. As shown in [link], the menses phase occurs during the 


early days of the follicular phase of the ovarian cycle, when progesterone, 
FSH, and LH levels are low. Recall that progesterone concentrations 
decline as a result of the degradation of the corpus luteum, marking the end 
of the luteal phase. This decline in progesterone triggers the shedding of the 
stratum functionalis of the endometrium. 

Hormone Levels in Ovarian and Menstrual Cycles 


Primordial Primary Secondary 
follicles follicles _ follicles 


= @Q=©)—_—_—- @ 


Atresia 
Single, selected 
om © —o-—— tertiary follicle 


i@) ==» © =» © —— & 
Atresia 
Constant development Selection of one dominant 
of early-stage follicles secondary follicle begins 
(2 months) each new menstrual cycle 


Ovarian cycle phases 


Selected tertiary Ovulation Corpus Corpus Degrading 
follicle luteum albicans corpus 


A g 
QO AN 6, B s 
0 7 14 21 28 
Day of menstrual cycle 


Uterine cycle phases 


0 ¥ 14 21 28 
Day of menstrual cycle 
Pituitary Ovulation ——FSH 
hormone levels ——LH 
5 0 7 14 21 28 
2 
a Ovarian —— Estrogen 
hormone levels === Progesterone 
0 rf 14 21 28 


Day of menstrual cycle 


The correlation of the hormone levels and their effects on 
the female reproductive system is shown in this timeline 
of the ovarian and menstrual cycles. The menstrual cycle 
begins at day one with the start of menses. Ovulation 
occurs around day 14 of a 28-day cycle, triggered by the 
LH surge. 


Proliferative Phase 


Once menstrual flow ceases, the endometrium begins to proliferate again, 
marking the beginning of the proliferative phase of the menstrual cycle 
(see [link]). It occurs when the granulosa and theca cells of the tertiary 
follicles begin to produce increased amounts of estrogen. These rising 
estrogen concentrations stimulate the endometrial lining to rebuild. 


Recall that the high estrogen concentrations will eventually lead to a 
decrease in FSH as a result of negative feedback, resulting in atresia of all 
but one of the developing tertiary follicles. The switch to positive feedback 
—which occurs with the elevated estrogen production from the dominant 
follicle—then stimulates the LH surge that will trigger ovulation. In a 
typical 28-day menstrual cycle, ovulation occurs on day 14. Ovulation 
marks the end of the proliferative phase as well as the end of the follicular 
phase. 


Secretory Phase 


In addition to prompting the LH surge, high estrogen levels increase the 
uterine tube contractions that facilitate the pick-up and transfer of the 
ovulated oocyte. High estrogen levels also slightly decrease the acidity of 
the vagina, making it more hospitable to sperm. In the ovary, the 
luteinization of the granulosa cells of the collapsed follicle forms the 


progesterone-producing corpus luteum, marking the beginning of the luteal 
phase of the ovarian cycle. In the uterus, progesterone from the corpus 
luteum begins the secretory phase of the menstrual cycle, in which the 
endometrial lining prepares for implantation (see [link]). Over the next 10 
to 12 days, the endometrial glands secrete a fluid rich in glycogen. If 
fertilization has occurred, this fluid will nourish the ball of cells now 
developing from the zygote. At the same time, the spiral arteries develop to 
provide blood to the thickened stratum functionalis. 


If no pregnancy occurs within approximately 10 to 12 days, the corpus 
luteum will degrade into the corpus albicans. Levels of both estrogen and 
progesterone will fall, and the endometrium will grow thinner. 
Prostaglandins will be secreted that cause constriction of the spiral arteries, 
reducing oxygen supply. The endometrial tissue will die, resulting in 
menses—or the first day of the next cycle. 


Note: 

Disorders of the... Feature 

Female Reproductive System 

Research over many years has confirmed that cervical cancer is most often 
caused by a sexually transmitted infection with human papillomavirus 
(HPV). There are over 100 related viruses in the HPV family, and the 
characteristics of each strain determine the outcome of the infection. In all 
cases, the virus enters body cells and uses its own genetic material to take 
over the host cell’s metabolic machinery and produce more virus particles. 
HPV infections are common in both men and women. Indeed, a recent 
study determined that 42.5 percent of females had HPV at the time of 
testing. These women ranged in age from 14 to 59 years and differed in 
race, ethnicity, and number of sexual partners. Of note, the prevalence of 
HPV infection was 53.8 percent among women aged 20 to 24 years, the 
age group with the highest infection rate. 

HPV strains are classified as high or low risk according to their potential to 
cause cancer. Though most HPV infections do not cause disease, the 
disruption of normal cellular functions in the low-risk forms of HPV can 
cause the male or female human host to develop genital warts. Often, the 


body is able to clear an HPV infection by normal immune responses within 
2 years. However, the more serious, high-risk infection by certain types of 
HPV can result in cancer of the cervix ([link]). Infection with either of the 
cancer-causing variants HPV 16 or HPV 18 has been linked to more than 
70 percent of all cervical cancer diagnoses. Although even these high-risk 
HPV strains can be cleared from the body over time, infections persist in 
some individuals. If this happens, the HPV infection can influence the cells 
of the cervix to develop precancerous changes. 

Risk factors for cervical cancer include having unprotected sex; having 
multiple sexual partners; a first sexual experience at a younger age, when 
the cells of the cervix are not fully mature; failure to receive the HPV 
vaccine; a compromised immune system; and smoking. The risk of 
developing cervical cancer is doubled with cigarette smoking. 
Development of Cervical Cancer 


@) During the G2 @® Cervix celis 
checkpoint, a with mutated remains 
functional p53 DNA do not healthy 


protein detects a divide 
DNA mutation 


HPV not 
present 


() p53 protein is With p53 Mutated cervix 
deactivated by the deactivated, cervix cells grow 

p53 inhibitor cells with mutated uncontrollably 
DNA successfully into a tumor 


In most cases, cells infected with the HPV virus heal on their 
own. In some cases, however, the virus continues to spread and 
becomes an invasive cancer. 


When the high-risk types of HPV enter a cell, two viral proteins are used to 
neutralize proteins that the host cells use as checkpoints in the cell cycle. 
The best studied of these proteins is p53. In a normal cell, p53 detects 
DNA damage in the cell’s genome and either halts the progression of the 


cell cycle—allowing time for DNA repair to occur—or initiates apoptosis. 
Both of these processes prevent the accumulation of mutations in a cell’s 
genome. High-risk HPV can neutralize p53, keeping the cell in a state in 
which fast growth is possible and impairing apoptosis, allowing mutations 
to accumulate in the cellular DNA. 

The prevalence of cervical cancer in the United States is very low because 
of regular screening exams called pap smears. Pap smears sample cells of 
the cervix, allowing the detection of abnormal cells. If pre-cancerous cells 
are detected, there are several highly effective techniques that are currently 
in use to remove them before they pose a danger. However, women in 
developing countries often do not have access to regular pap smears. As a 
result, these women account for as many as 80 percent of the cases of 
cervical cancer worldwide. 

In 2006, the first vaccine against the high-risk types of HPV was approved. 
There are now two HPV vaccines available: Gardasil® and Cervarix®. 
Whereas these vaccines were initially only targeted for women, because 
HPV is sexually transmitted, both men and women require vaccination for 
this approach to achieve its maximum efficacy. A recent study suggests 
that the HPV vaccine has cut the rates of HPV infection by the four 
targeted strains at least in half. Unfortunately, the high cost of 
manufacturing the vaccine is currently limiting access to many women 
worldwide. 


The Breasts 


Whereas the breasts are located far from the other female reproductive 
organs, they are considered accessory organs of the female reproductive 
system. The function of the breasts is to supply milk to an infant in a 
process called lactation. The external features of the breast include a nipple 
surrounded by a pigmented areola ((link]), whose coloration may deepen 
during pregnancy. The areola is typically circular and can vary in size from 
25 to 100 mm in diameter. The areolar region is characterized by small, 
raised areolar glands that secrete lubricating fluid during lactation to protect 
the nipple from chafing. When a baby nurses, or draws milk from the 
breast, the entire areolar region is taken into the mouth. 


Breast milk is produced by the mammary glands, which are modified 
sweat glands. The milk itself exits the breast through the nipple via 15 to 20 
lactiferous ducts that open on the surface of the nipple. These lactiferous 
ducts each extend to a lactiferous sinus that connects to a glandular lobe 
within the breast itself that contains groups of milk-secreting cells in 
clusters called alveoli (see [link]). The clusters can change in size 
depending on the amount of milk in the alveolar lumen. Once milk is made 
in the alveoli, stimulated myoepithelial cells that surround the alveoli 
contract to push the milk to the lactiferous sinuses. From here, the baby can 
draw milk through the lactiferous ducts by suckling. The lobes themselves 
are surrounded by fat tissue, which determines the size of the breast; breast 
size differs between individuals and does not affect the amount of milk 
produced. Supporting the breasts are multiple bands of connective tissue 
called suspensory ligaments that connect the breast tissue to the dermis of 
the overlying skin. 

Anatomy of the Breast 


Areolar glands 


Lactiferous 
Suspensory : sinuses 
ligament 


Nipple 


During lactation, milk moves from the alveoli 
through the lactiferous ducts to the nipple. 


During the normal hormonal fluctuations in the menstrual cycle, breast 
tissue responds to changing levels of estrogen and progesterone, which can 
lead to swelling and breast tenderness in some individuals, especially 
during the secretory phase. If pregnancy occurs, the increase in hormones 


leads to further development of the mammary tissue and enlargement of the 
breasts. 


Hormonal Birth Control 


Birth control pills take advantage of the negative feedback system that 
regulates the ovarian and menstrual cycles to stop ovulation and prevent 
pregnancy. Typically they work by providing a constant level of both 
estrogen and progesterone, which negatively feeds back onto the 
hypothalamus and pituitary, thus preventing the release of FSH and LH. 
Without FSH, the follicles do not mature, and without the LH surge, 
ovulation does not occur. Although the estrogen in birth control pills does 
stimulate some thickening of the endometrial wall, it is reduced compared 
with a normal cycle and is less likely to support implantation. 


Some birth control pills contain 21 active pills containing hormones, and 7 
inactive pills (placebos). The decline in hormones during the week that the 
woman takes the placebo pills triggers menses, although it is typically 
lighter than a normal menstrual flow because of the reduced endometrial 
thickening. Newer types of birth control pills have been developed that 
deliver low-dose estrogens and progesterone for the entire cycle (these are 
meant to be taken 365 days a year), and menses never occurs. While some 
women prefer to have the proof of a lack of pregnancy that a monthly 
period provides, menstruation every 28 days is not required for health 
reasons, and there are no reported adverse effects of not having a menstrual 
period in an otherwise healthy individual. 


Because birth control pills function by providing constant estrogen and 
progesterone levels and disrupting negative feedback, skipping even just 
one or two pills at certain points of the cycle (or even being several hours 
late taking the pill) can lead to an increase in FSH and LH and result in 
ovulation. It is important, therefore, that the woman follow the directions on 
the birth control pill package to successfully prevent pregnancy. 


Note: 


Aging and the... Feature 

Female Reproductive System 

Female fertility (the ability to conceive) peaks when women are in their 
twenties, and is slowly reduced until a women reaches 35 years of age. 
After that time, fertility declines more rapidly, until it ends completely at 
the end of menopause. Menopause is the cessation of the menstrual cycle 
that occurs as a result of the loss of ovarian follicles and the hormones that 
they produce. A woman is considered to have completed menopause if she 
has not menstruated in a full year. After that point, she is considered 
postmenopausal. The average age for this change is consistent worldwide 
at between 50 and 52 years of age, but it can normally occur in a woman’s 
forties, or later in her fifties. Poor health, including smoking, can lead to 
earlier loss of fertility and earlier menopause. 

As a woman reaches the age of menopause, depletion of the number of 
viable follicles in the ovaries due to atresia affects the hormonal regulation 
of the menstrual cycle. During the years leading up to menopause, there is 
a decrease in the levels of the hormone inhibin, which normally 
participates in a negative feedback loop to the pituitary to control the 
production of FSH. The menopausal decrease in inhibin leads to an 
increase in FSH. The presence of FSH stimulates more follicles to grow 
and secrete estrogen. Because small, secondary follicles also respond to 
increases in FSH levels, larger numbers of follicles are stimulated to grow; 
however, most undergo atresia and die. Eventually, this process leads to the 
depletion of all follicles in the ovaries, and the production of estrogen falls 
off dramatically. It is primarily the lack of estrogens that leads to the 
symptoms of menopause. 

The earliest changes occur during the menopausal transition, often referred 
to as peri-menopause, when a women’s cycle becomes irregular but does 
not stop entirely. Although the levels of estrogen are still nearly the same 
as before the transition, the level of progesterone produced by the corpus 
luteum is reduced. This decline in progesterone can lead to abnormal 
growth, or hyperplasia, of the endometrium. This condition is a concern 
because it increases the risk of developing endometrial cancer. Two 
harmless conditions that can develop during the transition are uterine 
fibroids, which are benign masses of cells, and irregular bleeding. As 
estrogen levels change, other symptoms that occur are hot flashes and night 
sweats, trouble sleeping, vaginal dryness, mood swings, difficulty 


focusing, and thinning of hair on the head along with the growth of more 
hair on the face. Depending on the individual, these symptoms can be 
entirely absent, moderate, or severe. 

After menopause, lower amounts of estrogens can lead to other changes. 
Cardiovascular disease becomes as prevalent in women as in men, possibly 
because estrogens reduce the amount of cholesterol in the blood vessels. 
When estrogen is lacking, many women find that they suddenly have 
problems with high cholesterol and the cardiovascular issues that 
accompany it. Osteoporosis is another problem because bone density 
decreases rapidly in the first years after menopause. The reduction in bone 
density leads to a higher incidence of fractures. 

Hormone therapy (HT), which employs medication (synthetic estrogens 
and progestins) to increase estrogen and progestin levels, can alleviate 
some of the symptoms of menopause. In 2002, the Women’s Health 
Initiative began a study to observe women for the long-term outcomes of 
hormone replacement therapy over 8.5 years. However, the study was 
prematurely terminated after 5.2 years because of evidence of a higher than 
normal risk of breast cancer in patients taking estrogen-only HT. The 
potential positive effects on cardiovascular disease were also not realized 
in the estrogen-only patients. The results of other hormone replacement 
studies over the last 50 years, including a 2012 study that followed over 
1,000 menopausal women for 10 years, have shown cardiovascular 
benefits from estrogen and no increased risk for cancer. Some researchers 
believe that the age group tested in the 2002 trial may have been too old to 
benefit from the therapy, thus skewing the results. In the meantime, intense 
debate and study of the benefits and risks of replacement therapy is 
ongoing. Current guidelines approve HT for the reduction of hot flashes or 
flushes, but this treatment is generally only considered when women first 
start showing signs of menopausal changes, is used in the lowest dose 
possible for the shortest time possible (5 years or less), and it is suggested 
that women on HT have regular pelvic and breast exams. 


Chapter Review 


The external female genitalia are collectively called the vulva. The vagina 
is the pathway into and out of the uterus. The man’s penis is inserted into 
the vagina to deliver sperm, and the baby exits the uterus through the 
vagina during childbirth. 


The ovaries produce oocytes, the female gametes, in a process called 
oogenesis. As with spermatogenesis, meiosis produces the haploid gamete 
(in this case, an ovum); however, it is completed only in an oocyte that has 
been penetrated by a sperm. In the ovary, an oocyte surrounded by 
supporting cells is called a follicle. In folliculogenesis, primordial follicles 
develop into primary, secondary, and tertiary follicles. Early tertiary 
follicles with their fluid-filled antrum will be stimulated by an increase in 
FSH, a gonadotropin produced by the anterior pituitary, to grow in the 28- 
day ovarian cycle. Supporting granulosa and theca cells in the growing 
follicles produce estrogens, until the level of estrogen in the bloodstream is 
high enough that it triggers negative feedback at the hypothalamus and 
pituitary. This results in a reduction of FSH and LH, and most tertiary 
follicles in the ovary undergo atresia (they die). One follicle, usually the one 
with the most FSH receptors, survives this period and is now called the 
dominant follicle. The dominant follicle produces more estrogen, triggering 
positive feedback and the LH surge that will induce ovulation. Following 
ovulation, the granulosa cells of the empty follicle luteinize and transform 
into the progesterone-producing corpus luteum. The ovulated oocyte with 
its surrounding granulosa cells is picked up by the infundibulum of the 
uterine tube, and beating cilia help to transport it through the tube toward 
the uterus. Fertilization occurs within the uterine tube, and the final stage of 
meiosis is completed. 


The uterus has three regions: the fundus, the body, and the cervix. It has 
three layers: the outer perimetrium, the muscular myometrium, and the 
inner endometrium. The endometrium responds to estrogen released by the 
follicles during the menstrual cycle and grows thicker with an increase in 
blood vessels in preparation for pregnancy. If the egg is not fertilized, no 
signal is sent to extend the life of the corpus luteum, and it degrades, 
stopping progesterone production. This decline in progesterone results in 
the sloughing of the inner portion of the endometrium in a process called 
menses, or menstruation. 


The breasts are accessory sexual organs that are utilized after the birth of a 
child to produce milk in a process called lactation. Birth control pills 
provide constant levels of estrogen and progesterone to negatively feed 
back on the hypothalamus and pituitary, and suppress the release of FSH 
and LH, which inhibits ovulation and prevents pregnancy. 


Interactive Link Questions 


Exercise: 
Problem: 
Watch this video to observe ovulation and its initiation in response to 


the release of FSH and LH from the pituitary gland. What specialized 
structures help guide the oocyte from the ovary into the uterine tube? 


Solution: 


The fimbriae sweep the oocyte into the uterine tube. 
Exercise: 
Problem: 
Watch this series of videos to look at the movement of the oocyte 
through the ovary. The cilia in the uterine tube promote movement of 


the oocyte. What would likely occur if the cilia were paralyzed at the 
time of ovulation? 


Solution: 


The oocyte may not enter the tube and may enter the pelvic cavity. 


Review Questions 


Exercise: 


Problem: What are the female gonads called? 


a. oocytes 
b. ova 

c. oviducts 
d. ovaries 


Solution: 


d 


Exercise: 


Problem: When do the oogonia undergo mitosis? 


a. before birth 

b. at puberty 

c. at the beginning of each menstrual cycle 
d. during fertilization 


Solution: 
a 


Exercise: 


Problem: From what structure does the corpus luteum originate? 


a. uterine corpus 

b. dominant follicle 
c. fallopian tube 

d. corpus albicans 


Solution: 


b 


Exercise: 


Problem: 
Where does fertilization of the egg by the sperm typically occur? 


a. vagina 
b. uterus 
c. uterine tube 
d. ovary 


Solution: 


C 


Exercise: 


Problem: Why do estrogen levels fall after menopause? 


a. The ovaries degrade. 

b. There are no follicles left to produce estrogen. 

c. The pituitary secretes a menopause-specific hormone. 
d. The cells of the endometrium degenerate. 


Solution: 


b 


Exercise: 


Problem: The vulva includes the 


a. lactiferous duct, rugae, and hymen 

b. lactiferous duct, endometrium, and bulbourethral glands 
c. mons pubis, endometrium, and hymen 

d. mons pubis, labia majora, and Bartholin’s glands 


Solution: 


d 


Critical Thinking Questions 


Exercise: 
Problem: 
Follow the path of ejaculated sperm from the vagina to the oocyte. 


Include all structures of the female reproductive tract that the sperm 
must swim through to reach the egg. 


Solution: 


The sperm must swim upward in the vagina, through the cervix, and 
then through the body of the uterus to one or the other of the two 
uterine tubes. Fertilization generally occurs in the uterine tube. 


Exercise: 


Problem: 


Identify some differences between meiosis in men and women. 


Solution: 


Meiosis in the man results in four viable haploid sperm, whereas 
meiosis in the woman results in a secondary oocyte and, upon 
completion following fertilization by a sperm, one viable haploid 
ovum with abundant cytoplasm and up to three polar bodies with little 
cytoplasm that are destined to die. 


Exercise: 


Problem: 


Explain the hormonal regulation of the phases of the menstrual cycle. 


Solution: 


As aresult of the degradation of the corpus luteum, a decline in 
progesterone concentrations triggers the shedding of the endometrial 
lining, marking the menses phase of the menstrual cycle. Low 
progesterone levels also reduce the negative feedback that had been 
occurring at the hypothalamus and pituitary, and result in the release of 
GnRH and, subsequently, FSH and LH. FSH stimulates tertiary 
follicles to grow and granulosa and theca cells begin to produce 
increased amounts of estrogen. High estrogen concentrations stimulate 
the endometrial lining to rebuild, marking the proliferative phase of the 
menstrual cycle. The high estrogen concentrations will eventually lead 
to a decrease in FSH because of negative feedback, resulting in atresia 
of all but one of the developing tertiary follicles. The switch to positive 
feedback that occurs with elevated estrogen production from the 
dominant follicle stimulates the LH surge that will trigger ovulation. 
The luteinization of the granulosa cells of the collapsed follicle forms 
the progesterone-producing corpus luteum. Progesterone from the 
corpus luteum causes the endometrium to prepare for implantation, in 
part by secreting nutrient-rich fluid. This marks the secretory phase of 
the menstrual cycle. Finally, in a non-fertile cycle, the corpus luteum 
will degrade and menses will occur. 


Exercise: 


Problem: 


Endometriosis is a disorder in which endometrial cells implant and 
proliferate outside of the uterus—in the uterine tubes, on the ovaries, 
or even in the pelvic cavity. Offer a theory as to why endometriosis 
increases a woman’s risk of infertility. 


Solution: 


Endometrial tissue proliferating outside of the endometrium—for 
example, in the uterine tubes, on the ovaries, or within the pelvic 
cavity—could block the passage of sperm, ovulated oocytes, or a 
zygote, thus reducing fertility. 


Glossary 


alveoli 
(of the breast) milk-secreting cells in the mammary gland 


ampulla 
(of the uterine tube) middle portion of the uterine tube in which 
fertilization often occurs 


antrum 
fluid-filled chamber that characterizes a mature tertiary (antral) follicle 


areola 
highly pigmented, circular area surrounding the raised nipple and 
containing areolar glands that secrete fluid important for lubrication 
during suckling 


Bartholin’s glands 
(also, greater vestibular glands) glands that produce a thick mucus that 
maintains moisture in the vulva area; also referred to as the greater 
vestibular glands 


body of uterus 
middle section of the uterus 


broad ligament 
wide ligament that supports the uterus by attaching laterally to both 
sides of the uterus and pelvic wall 


cervix 
elongate inferior end of the uterus where it connects to the vagina 


clitoris 
(also, glans clitoris) nerve-rich area of the vulva that contributes to 
sexual sensation during intercourse 


corpus albicans 


nonfunctional structure remaining in the ovarian stroma following 
structural and functional regression of the corpus luteum 


corpus luteum 
transformed follicle after ovulation that secretes progesterone 


endometrium 
inner lining of the uterus, part of which builds up during the secretory 
phase of the menstrual cycle and then sheds with menses 


fimbriae 
fingerlike projections on the distal uterine tubes 


follicle 
ovarian structure of one oocyte and surrounding granulosa (and later 
theca) cells 


folliculogenesis 
development of ovarian follicles from primordial to tertiary under the 
stimulation of gonadotropins 


fundus 
(of the uterus) domed portion of the uterus that is superior to the 
uterine tubes 


granulosa cells 
supportive cells in the ovarian follicle that produce estrogen 


hymen 
membrane that covers part of the opening of the vagina 


infundibulum 
(of the uterine tube) wide, distal portion of the uterine tube terminating 
in fimbriae 


isthmus 
narrow, medial portion of the uterine tube that joins the uterus 


labia majora 


hair-covered folds of skin located behind the mons pubis 


labia minora 
thin, pigmented, hairless flaps of skin located medial and deep to the 
labia majora 


lactiferous ducts 
ducts that connect the mammary glands to the nipple and allow for the 
transport of milk 


lactiferous sinus 
area of milk collection between alveoli and lactiferous duct 


mammary glands 
glands inside the breast that secrete milk 


menarche 
first menstruation in a pubertal female 


menses 
shedding of the inner portion of the endometrium out though the 
vagina; also referred to as menstruation 


menses phase 
phase of the menstrual cycle in which the endometrial lining is shed 


menstrual cycle 
approximately 28-day cycle of changes in the uterus consisting of a 
menses phase, a proliferative phase, and a secretory phase 


mons pubis 
mound of fatty tissue located at the front of the vulva 


myometrium 
smooth muscle layer of uterus that allows for uterine contractions 
during labor and expulsion of menstrual blood 


oocyte 


cell that results from the division of the oogonium and undergoes 
meiosis I at the LH surge and meiosis II at fertilization to become a 
haploid ovum 


oogenesis 
process by which oogonia divide by mitosis to primary oocytes, which 
undergo meiosis to produce the secondary oocyte and, upon 
fertilization, the ovum 


oogonia 
ovarian stem cells that undergo mitosis during female fetal 
development to form primary oocytes 


ovarian cycle 
approximately 28-day cycle of changes in the ovary consisting of a 
follicular phase and a luteal phase 


ovaries 
female gonads that produce oocytes and sex steroid hormones (notably 
estrogen and progesterone) 


ovulation 
release of a secondary oocyte and associated granulosa cells from an 
Ovary 


ovum 
haploid female gamete resulting from completion of meiosis II at 
fertilization 


perimetrium 
outer epithelial layer of uterine wall 


polar body 
smaller cell produced during the process of meiosis in oogenesis 


primary follicles 
ovarian follicles with a primary oocyte and one layer of cuboidal 
granulosa cells 


primordial follicles 
least developed ovarian follicles that consist of a single oocyte and a 
single layer of flat (squamous) granulosa cells 


proliferative phase 
phase of the menstrual cycle in which the endometrium proliferates 


rugae 
(of the vagina) folds of skin in the vagina that allow it to stretch during 
intercourse and childbirth 


secondary follicles 
ovarian follicles with a primary oocyte and multiple layers of 
granulosa cells 


secretory phase 
phase of the menstrual cycle in which the endometrium secretes a 
nutrient-rich fluid in preparation for implantation of an embryo 


suspensory ligaments 
bands of connective tissue that suspend the breast onto the chest wall 
by attachment to the overlying dermis 


tertiary follicles 
(also, antral follicles) ovarian follicles with a primary or secondary 
oocyte, multiple layers of granulosa cells, and a fully formed antrum 


theca cells 
estrogen-producing cells in a maturing ovarian follicle 


uterine tubes 
(also, fallopian tubes or oviducts) ducts that facilitate transport of an 
ovulated oocyte to the uterus 


uterus 
muscular hollow organ in which a fertilized egg develops into a fetus 


vagina 


tunnel-like organ that provides access to the uterus for the insertion of 
semen and from the uterus for the birth of a baby 


vulva 
external female genitalia